Migrations and Management of the Jackson Elk Herd

19970319 030

U.S. DEPARTMENT OF THE INTERIOR NATIONAL BIOLOGICAL SURVEY I Resource Publication 199 Technical Report Series National Biological Survey

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By Bruce L. Smith Russell L. Robbins

U.S. DEPARTMENT OF THE INTERIOR NATIONAL BIOLOGICAL SURVEY Resource Publication 199 Washington, D.C. • 1994 Contents Page Foreword . V Abstract .. 1 Objectives . 7 Study Area 7 Methods .. 9 Definitions 9 Radio Performance 10 Monitoring with Radiotelemetry 10 Delineation of Seasonal Ranges 11 Delineation of Calving Areas . . 12 Estimated Sizes and Age-Class Compositions of Herd Segments . 12 Evaluation of Harvests 12 Statistical Treatment 13 Results ...... 13 Summer Distribution . 13 Fidelity to Winter Ranges . 13 Fidelity to Summer Ranges . 14 Spring Migration 14 Fall Migration 17 Calving. 23 Natality .. . 28 Harvests .. . 30 Natural Mortality 32 Survival .. 34 Discussion . . 34 Distribution 34 Migration . 35 Calving ... 37 Sympatry and Diseases of Wildlife and Livestock . 38 Survival ...... 41 Harvests of Elk and Population Regulation . . . . 41 Size and Composition of the Grand Teton National Park Herd Segment 44 Population Regulation ...... 45 Restoration of Historic Distributions of Elk 46 Recommended Research . . . 48 Recommended Management . 48 Population Management 48 Brucellosis Management 50 Acknowledgments 50 Cited Literature 50 Appendix A 54 AppendixB 55 AppendixC 56 AppendixD 61 iii Foreword The Jackson elk herd is one of the large elk herds in the Greater Yellowstone Ecosystem and has gained international recognition and caused considerable debate. The recognition is based on the long tradition ofresearch that began in 1927 with the assignment of Olaus Murie to the National Elk Refuge for the explicit purpose of conducting research on elk. Much of the data Murie collected was used in his book The elk of North America, the first comprehensive book on that topic. Russell "Buzz" Robbins assumed Murie's position of biologist with the National Elk Refuge in 1968. He conducted research on methods for improving the elk winter feeding program, the topic of considerable debate. A natural extension of that feeding program was to determine more precisely where the elk that winter on the National Elk Refuge spend summer and fall and how the timing of their migrations affect the harvest of these elk in fall. When Robbins assumed a position with the Arctic National Wildlife Refuge, Bruce Smith became refuge biologist. Since 1982, Smith has pursued research on the Jackson elk herd with candor and a special understanding of this large ungulate and its value to the various segments of the public. The Migrations and Management of the Jackson Elk Herd presents the authors' 25-year collection of data that include one of the most comprehensive sets ofavailable data on ungulate migrations. The authors integrated their own data not only with the literature but with the numerous management reports of the past 40 years. The work is an important and comprehensive analysis of the problems and management of one of the most controversial herds of ungulates in North America.

John L. Oldemeyer Leader Endangered Species Research National Ecology Research Center National Biological Survey

V Migrations and Management of the Jackson Elk Herd

by

Bruce L. Smith and Russell L. Robbins1

U.S. Fish and Wildlife Service National Elk Refuge P.O. BoxC Jackson, Wyoming 83001

Abstract. From April 1978 to April 1982, 85 adult (~2. 7 years-old) elk (Cervus elaphus nelsoni) were captured and radio-collared on the National Elk Refuge in northwestern Wyoming. Relocations with biotelemetry from 1978 to 1984 showed that the radio-collared elk occupied four distinct summer ranges: 48% summered in Grand Teton National Park, 28% in Yellowstone National Park, 12% in the Teton Wilderness, and 12% in the Gros Ventre drainage. Elk that summered in a specific range are collectively a herd segment of the Jackson elk herd. The fidelity to the summer ranges was 98%; the fidelity to the winter range at the National Elk Refuge was 97%. At least 69% of the calving by elk from all four summer ranges occurred in the Grand Teton National Park central valley. The fidelity to calving areas inversely correlated with the distance between a calving area and the National Elk Refuge winter range. The fall migration commenced in October or November, and 94% of the radio-collared elk arrived at the National Elk Refuge by 15 December. Most elk that were harvested by hunters in the park and on the National Elk Refuge during the early stages of the fall migration were from the Grand Teton National Park herd segment. From 1978 to 1984, hunters reduced the number of wintering elk on the National Elk Refuge from 8,413 to 5,010. Efforts to reduce the size of the Grand Teton National Park herd segment by hunting have been unsuccessful. However, the size of this herd segment seemingly stabilized since the late 1960s and was about 3,700 elk (2,869-4,486, 95% CI, post-hunting season) during this study.

We recommend the establishment of management objectives for each of the four herd segments and an investigation of density-dependent effects on the recruitment, survival, and dispersal ofjuveniles. Reducing the number of elk in Grand Teton National Park and restoring the number and migrations of elk east of Grand Teton National Park to pre-1950 conditions are goals of the state and federal agencies that manage the elk and their habitat. Alternative management based on this research is offered to accomplish those goals.

Key words: Brucellosis, Cervus elaphus, elk, harvest, management, migration, survival, Wyoming.

1 Present address: Route 2, Box 50, Elk Point, South Dakota 57025. 1 2 RESOURCE PUBLICATION 199

The management of the Jackson elk (Cervus herd winters and is supplementally fed at the Na­ elaphus) herd in northwestern Wyoming is a com­ tional Elk Refuge and at three state-operated feed plex interagency effort. The elk of the Jackson elk grounds in the Gros Ventre River drainage (Fig. 2). herd are known for their winter concentrations and The U.S. Fish and Wildlife Service and the extensive annual migrations (as long as 90 km) Wyoming Game and Fish Department work un­ between their winter and summer ranges. They der a 197 4 cooperative agreement to manage the migrate through and summer in the Bridger-Teton National Elk Refuge for a maximum of 7,500 elk, National Forest, Yellowstone National Park, Grand which is 68% of the targeted post-hunting-season Teton National Park, and scattered parcels of pri­ size of the entire Jackson elk herd. vate and state lands (Fig. 1). Some elk winter in the By the early 1900s, fewer than 50,000 elk re­ Bridger-Teton National Forest and Grand Teton mained in North America. Most were concen­ National Park, but about 90% of the Jackson elk trated in the area of Yellowstone National Park

A 9-month-old elk eating alfalfa pel­ lets on the National Elk Refuge, Wyoming; March 1991. MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 3

,..._.....," .... -- Park boundaries ', - Roads ', """ -- Rivers , } I • - • - Nat. Elk Refuge <: I } ~ ~ 1 0246810- L_t\ Yellowstone Kilometers '"-'--1.,National I ---:_;--T- Park I Memoria, Parkway ~ .J Teton /., l_ Wilderness < j Grand Teton National Park 'l \ ) I \

Gros Ventre Range

Fig. 1. The northern Jackson Hole region. 4 RESOURCE PUBLICATION 199

....., -- Park boundaries .... I 8 8 Gros Ventre feedgrounds • • • • Herd segments I YNP •• I ~-~ Yellowstone 0246810 1 Kilometers National . .. Park : I ------~­...... •":'I· ,:-· L·. . . /." - ~ <: ~· TW 1: GTNP f5 ~ } .. ~ . . ... \ .. . L:. ...:..: :.:. .-i": % \ . F ;:, . . ( I: •• Buffalo Fork River .. - ... . l . ... \ Grand ...... J Teton .,,-- ...... J• ...... • • • •r.• I • I ~ational J:·· GV .. . \P.ark 1-·· ... ~,: ...... Fig. 2. Summer distributions and winter feed grounds of the four herd segments of the Jackson elk herd: Grand Teton National Park (GTNP), Yellowstone National Park (YNP), Teton Wilderness (TW), and Gros Ventre drainage (GV). MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 5

and Jackson Hole (Seton 1927). Accounts of the winter was on an emergency basis until the abundance of elk in Jackson Hole prior to Euro­ 1950s when annual feeding operations began (An­ pean settlement in 1884 are limited to observa­ derson 1958). tions by fur trappers and explorers (Anderson After establishment of the National Elk Refuge, 1958; Robbins et al. 1982). Postsettlement esti­ the number of elk in Jackson Hole began to stabi­ mates and censuses of elk wintering in the Jack­ lize, although the number and probably the distri­ son Hole valley were made by game wardens and bution of elk in what is now Grand Teton National U.S. Forest Service personnel. Cole (1969:15) Park apparently declined between 1900 and 1950 stated: (National Park Service. 1986. Grand Teton Na­ tional Park Natural Resources Management Plan The 1887 to 1911 estimates of 15,000 to and Environmental Assessment. Moose, Wyo. 25,000 elk in the Jackson Hole herd, with the largest numbers reported during severe 459 pp.). Established in 1929, Grand Teton Na­ winters, should establish that the Jackson tional Park originally included only the Teton Hole and Gros Ventre Valleys were historical Range. Cattle ranching and dude ranches (to serve winter areas. The first organized censuses a growing tourist industry) occupied the northern (beginning in 1912 and covering Jackson Hole Jackson Hole Valley (lands north of the National and the Hoback River drainage south of Elk Refuge between the Teton Range on the west Jackson), which accounted for approximately and the Gros Ventre Mountains and mountains of 20,000 animals, also seem to confirm that the Teton Wilderness on the north and east), which large numbers of elk were present was added to Grand Teton National Park in 1950. historically. This addition to the park is known as the central valley of Grand Teton National Park. Prior to This statement disputes claims that elk began 1950, fewer elk than now summered in the central wintering in Jackson Hole only after settlement valley (Anderson 1958; Cole 1969). blocked migration routes from the valley to his­ The state of Wyoming resisted expansion of toric winter ranges to the south (Anderson 1958). Grand Teton National Park for fear that elk pro­ In 1911, the Congress authorized an investiga­ tected in the park would increase and predomi­ tion of the status of the elk herds in Jackson Hole nate among the elk wintering on the National Elk (Preble 1911). Preble estimated that 20,000 elk Refuge and among the elk on the feed grounds in summered north of Jackson: 8,000 between the the Gros Ventre drainage (Blunt 1950) and there­ Gros Ventre and Buffalo Fork rivers and another fore in the Jackson elk herd. Consequently the 12,000 north of the Buffalo Fork River of which no legislation (Public Law 81-787) specified that re­ more than 4,000 were in Yellowstone National ductions of elk be permitted in designated portions Park (Fig. 1). of the expanded Grand Teton National Park. The establishment of Yellowstone National The public's interest and demand for elk in­ Park in 1872 and the National Elk Refuge in 1912 were landmarks in the protection of elk from mar­ creased after 1950 when competing uses of elk ket and tusk hunting and usurpation of critical habitat on public and private lands for timber and habitat for human uses. In 1912, the Congress energy production, recreation, living space for the established the National Elk Refuge as a winter human population of Jackson Hole, and visitation range for elk in Jackson Hole. Feeding of elk in to the area grew (C. Phillips and S. Ferguson. winter was initiated to mitigate losses of historic 1977. Hunting and fishing expenditure values and winter ranges after the settlement of Jackson Hole participation preferences in Wyoming, 1975. Uni­ by Europeans in 1884 and attendant livestock versity of Wyoming, Laramie, unpublished; Rippe grazing and haying on winter ranges. Sub­ and Rayburn 1981; Roelle and Auble 1983). Inten­ sequent conflicts between elk and ranchers and sified resource development in the Bridger-Teton periodic heavy mortality of elk during severe win­ National Forest, expanding road systems, upgrad­ ters typified the 1890s and early 1900s (Brown ing of existing roads, increased hunting pressure, 1947; Anderson 1958; Robbins et al. 1982). Feed­ loss of winter range by development of private ing of the elk in the Gros Ventre drainage during lands, and expansion of Grand Teton National 6 RESOURCE PUBLICATION 199

Park in 1950 altered the distribution and harvest His methods included (1) repeated ground and of the elk. aerial counts on sample areas; (2) recording of the Hunting of the Jackson elk herd has long been proportion of elk that had been neck-banded on an attraction and a source of debate (C. Sheldon the National Elk Refuge in winter and had been 1927. The conservation of the elk of Jackson Hole, observed on various summer ranges; and (3) track Wyoming: A report to the President's committee on counts of migrating elk along a 76-km-road tran­ outdoor recreation and the Governor of Wyoming, sect and periodic counts of elk arriving at the unpublished; Brown 1947; Murie 1951). Some ob­ National Elk Refuge. The road transect started at servers perceive irony in the maintenance of a Moose, Wyoming, and extended north through the population of elk at high numbers by tax-subsi­ west side of the park to Moran, Wyoming, and then dized feeding and the hunting in a national park east to the top of the Togwotee Pass (Fig. 1). to control the size of the same population (Wood Cole's (1969) analysis and Boyce's (1989) up­ 1984). Elk hunting on the National Elk Refuge has dated analysis through 1985 illustrate that the likewise received adverse national attention (Dex­ proportion of elk that migrated through Grand ter 1984). However, the mortality of the Jackson Teton National Park from Yellowstone National elk herd in winter is low, and hunting on those Park and the Teton Wilderness drastically in­ federal lands has been important to controlling creased since 1950. Because the counts do not the herd's size (Thomas et al. 1984; Boyce 1989). include tracks of elk that migrate from the Gros Individuals who favor hunting disagree on where Ventre drainage to the National Elk Refuge or and how the elk should be harvested. Some tracks of all elk that summer in Grand Teton want even more elk harvested because of direct or National Park, the track counts cannot be used indirect economic interests (Taylor et al. 1982; alone to determine the proportions of elk in each Boyce 1989). herd segment. Furthermore, when they cross the Since the expansion of Grand Teton National road transect between Moran and the Togwotee Park, a growing concern of managers has been to Pass, elk of the Teton Wilderness herd segment improve the distribution of elk on summer ranges cannot be distinguished from elk of the Yellow­ (i.e., more elk on national forest lands and fewer stone National Park herd segment. Therefore, in Yellowstone National Park [Anderson 1958; Boyce (1989) combined the elk from the Yellow­ Cole 1969]). Anderson (1958) and later Cole (1969) stone National Park and Teton Wilderness herd advocated (1) increasing harvests of the Yellow­ segments in his analysis. Finally, varying num­ stone National Park herd segment that travelled bers of elk that cross the transects each year in the through the roadless Teton Wilderness before absence of snow do not leave tracks and are there­ crossing Grand Teton National Park because that fore not recorded. herd segment had become too large to manage and Test results of about 28% of the elk that winter (2) allowing compensatory increases in elk that on the National Elk Refuge (39% of the adult migrated through roaded areas east of Grand Te­ females) are positive for Brucella abortus antibod­ ton National Park or summered on the national ies (Boyce 1989). Brucellosis, which is caused by forest lands between the two parks. To the pre­ B. abortus, not only affects the health and produc­ sent, this has included an annual elk hunt in tivity of the Jackson elk herd (Thorne et al. 1991; Grand Teton National Park. Cole (1969) also urged Oldemeyer et al. 1993) but is potentially transmis­ hunting in Grand Teton National Park to control sible among cattle, bison, and elk (Thorne et al. the increasing number of elk that summered there 1982; Davis et al. 1988). The Jackson bison herd and wintered on the National Elk Refuge. The elk is also infected with brucellosis (Williams et al. that summered in Grand Teton National Park 1993). Because of the cattle ranching in Jackson were the earliest migrants to the National Elk Hole, an understanding of the spatial and tempo­ Refuge each fall and consumed forage intended for ral overlap of seasonal ranges of the three species winter grazing. and potential risk of brucellosis transmission is Cole (1969) estimated the proportion of the elk essential to management of the Jackson elk and that wintered on the refuge and summered on bison herds and to planning of grazing allotments various ranges in Jackson Hole during the 1960s. on federal lands. MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 7

Objectives Hole and the surrounding Teton Range to the west, Gros Ventre Range to the east, and moun­ tains of southern Yellowstone National Park and In 1978, we initiated this study of the approxi­ the Teton Wilderness to the north. The Fall Creek mately 7,500 elk that winter on the National Elk elk herd occupies the Snake River and lower Ho­ Refuge. We monitored elk with radiotelemetry dur­ back River drainages of southern Jackson Hole ing 1978-84 to evaluate management and sub­ south of the town of Jackson. The Jackson sequent changes in the elk population. During the elk herd's distribution (the Jackson herd unit) en­ study, the number of wintering elk on the refuge compasses 5,195 km2 in the Snake River water­ declined by 40%. We had 8 objectives. shed which drains to the south, and the Yellow­ 1. Determine the seasonal distributions and mi­ ston~ River watershed, which drains to the grations of elk that winter on the National Elk north (Fig. 1). Refuge. Elevations range from 1,890 m on the National 2. Evaluate Cole's (1969) prediction that the coop­ Elk Refuge to more than 3,600 min the mountains. erative management of the National Park Serv­ Geologic features were described by Love and Reed ice and Wyoming Game and Fish Department (1968) and Houston (1968); the vegetation was de­ will restore the historic (pre-1950) migrations scribed by Cole (1969). Plant communities include and distributions of the Jackson elk herd by riparian cottonwood (Populus angustifolia) wood­ reducing the Yellowstone National Park herd lands and sagebrush (Artemisia spp.) grasslands in segment and increasing the Teton Wilderness the Jackson Hole valley; aspen (Populus tremu­ herd segment. loides ), Douglas fir (Pseudotsuga menziesii), and 3. Identify calving locations and their relation to lodgepole pine (Pinus contorta) forests in the foot­ distributions of the Jackson elk herd. hills and on the lower mountain slopes; spruce 4. Identify migration routes and timing of migra­ (Picea engelmannii) and fir (Abies lasiocarpa) for­ tions. Fall-migration routes were described by ests and tall forb meadows in the subalpine zone; Anderson (1958) and Cole (1969), and their rela­ and arctic alpine vegetation above 3,050 m. tive use was estimated from track counts in the The average annual precipitation at Moran, snow (Boyce 1989). We sought to determine Wyoming, is 57 cm. The mean annual temperature which routes elk from the various summer is 1. 7° C and ranges from monthly means of-11.0° ranges used and if fall migrations from all sum­ C in January to 14.7° C in July (National Oceanic mer ranges were initiated by snow accumula­ and Atmospheric Administration 1982). Elk that tions (Rudd et al. 1983, Boyce 1991). 5. Determine the degree of fidelity of elk to winter summer in the Teton Wilderness and Gros Ventre ranges, herd segments, calving areas, and mi­ drainage occupy lands (primarily the Bridger-Te­ gration routes. ton National Forest) that are open to big-game 6. Estimate the annual harvest rates of elk in hunting in the fall. Yellowstone National Park is each herd segment. (Rudd et al. 1983, Boyce closed to hunting, but elk that summer there are 1991). subject to hunting as they migrate south from the 7. Estimate survival rates and determine whether park. Elk that summer in or migrate through survival rates of elk that summered inside and portions of Grand Teton National Park, in which outside Grand Teton National Park differed. hunting is permitted under Public Law 81-787 8. Identify where distributions of elk, cattle, and (the legislation for the expansion of the park in bison spatially or temporally overlap. 1950), can be hunted. Those areas are also desig­ nated Wyoming Hunting Units 75, 76, and 79 and are east of U.S. Highway 89/287 and the Snake River (Fig. 3). Hunting Unit 72 in the northwest Study Area corner of the park has been closed to hunting since 1967. A limited elk hunt is held on the northern The Jackson elk herd roams the northern por­ half of the National Elk Refuge, designated as tions of an intermountain basin known as Jackson State-Hunting Unit 77. 8 RESOURCE PUBLICATION 199

- Hunting unit boundaries -- Park boundaries • -• Nat Elk Refuge I Yellowstone 1 ~-~ National 0 2 4 6 8 10 Park 1 Kilometers ___ _J ------

~ C } \ Grand J Teton 1 National __,\Park \ J ("' r1.. ,J

Fig. 3. Hunting units in the Jackson elk herd unit. MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 9

Methods the same technique and replaced nonfunctioning radios and radios nearing the life expectancy of the batteries. Three of the 35 elk were recaptured Definitions twice (Appendix B). The sample of radio-collared elk is a small pro­ The geographical range of the Jackson elk herd portion of the elk that winter on the National Elk is 5,195 km2 in the Snake River and Yellowstone Refuge, and the randomness of the sample may be River watersheds and is known as the Jackson herd questioned. However, money and time constraints unit (Cole 1969; Fig. 1). In summer, the Jackson elk precluded a larger sample. We offer the following herd separates into four herd segments, each of arguments that the sample was adequate for mean­ which summers on one of four ranges: Grand Teton ingful inferences. National Park, Yellowstone National Park, the Te­ After their arrival on the National Elk Refuge ton Wilderness, and the Gros Ventre drainage each fall, elk from the four herd segments were (Fig. 2). The Grand Teton National Park summer assumed to mingle. The arrival of the elk coincided range is divided into Northwest Grand Teton Na­ with the elk hunt on the northern half of the refuge tional Park and Grand Teton National Park valley. and with early-winter snowfall. The elk congre­ Grand Teton National Park valley refers to all lands gated in one or two large groups on the southern south and east of Jackson Lake and comprises the one-third of the refuge until the conclusion of the summer range of most of the Grand Teton National hunt. From then until supplemental feeding com­ Park herd segment (Fig. 1). In winter, elk of the menced in January, the elk ranged freely in dy­ herd segments mingle on the National Elk Refuge namic groups. and on other ranges of the Jackson herd unit and We provided the elk with supplemental feed at 80-90% are provided with supplemental feed on four feed grounds that were separated from each feed grounds. other by 1-2 km. After each morning's feeding, the elk bedded until mid-afternoon. While grazing on Capture of Elk for Radio-Collaring the flat grasslands on the refuge during the evening During early April of the years 1978-82, 76 and night, elk from adjacent feed grounds mingled. pregnant, 2::2. 7 year-old elk were radio-collared No physical barriers inhibited the elk from moving and released. These females were obtained from among the four feed grounds. those used in feeding trials on the National Elk We trapped and immobilized elk at random. The Refuge to determine maintenance rations of win­ trap was near the center of the refuge where elk ter feed for elk (Oldemeyer et al. 1993). After the from all feed grounds on the refuge mingled. We elk were captured in a baited corral trap, they immobilized elk on two different feed grounds. We were placed in 2- or 4-ha pastures. Their ages were determined that proportions of radio-collared elk determined from tooth eruption and wear that migrated to each summer range were not dif­ (Quimby and Gaab 1957). Pregnancy was deter­ ferent (X2 = 1.97, df = 3, P = 0.58; Table 1). We mined by palpation of recta (Greer and Hawkins therefore combined the trapped and immobilized 1967). In 1978, 1979, and 1980, every other female elk in one sample for data analyses. elk was radio-collared and released during April Subsequent monitoring indicated that radio-col­ when all elk in the feeding trials were retrapped lard elk randomly mingled on the National Elk for weighing. In 1982 (trials were not conducted in Refuge. The frequency of daily feed-ground atten­ 1981), collars were placed on only six elk in April, dance by radio-collared elk was not different among one on every tenth animal (Appendix A). the three feed grounds on which elk were captured We captured and radio-collared another 21 2::2. 7 and radio-collared (F = 0.55; df = 2, 56; P = 0.58). year-old elk (17 males and 4 females) during Elk from all four herd segments freely traveled March and April of 1981 by immobilizing them between feed grounds during winter. with powdered succinylcholine chloride-loaded We also compared the expected with the ob­ Pneu-Darts (Pneu-Dart, 406 Bridge Street, Wil­ served use offeed grounds by radio-collared elk on liamsport, Pa. 17701) that we shot from trucks. We the refuge. The frequency of attendance, based on recaptured 35 elk on the National Elk Refuge with relocations with telemetry, was considered use, 10 RESOURCE PUBLICATION 199

Table 1. Numbers (n) and proportions (p) of elk (Ceruus elaphus) of herd segmentsa of the Jackson elk herd that were trapped and immobilized on the National Elk Refuge, Wyoming, 1978-1982. The proportion of trapped and immobilized elk did not differ between herd segments. Herd segment Trapped Immobilized Test result (summer range) n p n p z p Grand Teton National Park 29 0.45 12 0.57 -0.77 0.22 Yellowstone National Park 19 0.30 5 0.24 0.40 0.34 Teton Wilderness 7 0.11 3 0.14 -0.19 0.42 Gros Ventre Drainage 9 0.14 1 0.05 0.67 0.25 aThe herd segments are identified by their summer ranges.

and mean daily numbers of elk at each feed ground Embossed instructions on the radio collars re­ during the study was considered availability. Si­ quested hunters to return the collars to the Na­ multaneous 95% confidence intervals were con­ tional Elk Refuge and to provide information on structed around the proportions of use of the four the date and place of harvest. We retrieved collars feed grounds (Neu et al. 1974; Byers et al. 1984; of elk believed dead from natural causes. Ages at Table 2). Preference was not demonstrated (X2 = the time of death were determined from an analy­ 1.07, P = 0. 79). Based on these analyses, we be­ sis of the cementum annuli (Mitchell 1963). lieve that the distributions of the radio-collared elk on summer ranges provided an unbiased esti­ mator of the overall summer distribution of the elk Monitoring with Radiotelemetry that wintered on the refuge. We attempted to locate transmitters 3 times/week during migration and approximately Radio Performance weekly at other times by triangulation with mobile ground-based receiving equipment (Appendix A). Personnel of the Denver Wildlife Research Cen­ We transported the three-element yagi antennas, ter built the 24 radio collars placed on elk in 1978 AVM-164-MHz receivers, and headphones by vehi­ and another 34 placed on elk in 1979 (17 replaced cle and on foot and horseback to strategic signal-re­ nonfunctioning collars from 1978). Telonics, Inc. ceiving locations. During April-December, Cessna (Mesa, Arizona) manufactured the radio collars 182 and 206, Piper Super Cub, and Maule fixed­ placed on elk in 1980-82. Each elk was monitored wing aircraft were used to locate elk. Three-ele­ for ::;60 months (Appendix A). ment, unidirectional yagi antennas, attached to

0 Table 2. Confidence intervals of the observed (Pi) and expected (P / proportions of use of the four feed grounds on the National Elk Refuge, Wyoming, by radio-collared elk (Ceruus elaphus) of the Jackson elk herd, 1978-1984. All confidence intervals were constructed with the Bonferroni procedure and contained the expected proportions. Proportional use 95% Confidence

Feed ground Observed (pi) Expected (?i0 ) intervals Shop 0.113 0.130 0.046-0.179 Nowlin 0.310 0.300 0.213-0.407 Poverty Flats 0.296 0.317 0.200-0.391 McBride 0.282 0.252 0.187-0.376 aByers et al. 1984. MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 11

Grazing by livestock and haying of historic winter ranges of elk in Jackson Hole, Wyoming, led to severe mortality of elk during some winters and to the subsequent supplemental feeding of the Jackson elk herd. Photo by S.N. Leek [deceased] 1909. each wing strut and linked to a Telonics receiver­ (all lands in Grand Teton National Park west of scanner and right-left switch box, were used to Jackson Lake) and Grand Teton National Park locate elk that migrated beyond the range ofreceiv­ valley (the remaining southern and central part of ing equipment on the ground. Grand Teton National Park that is the summer range of most elk of the Grand Teton National Park herd segment). Fidelity to seasonal ranges was Delineation of Seasonal Ranges measured as an elk's frequency of return to its previous year's winter range and its previous year's Radio locations were plotted on USGS topo­ summer range. Elk were monitored for as many as graphic maps and recorded by the Universal Trans­ 5 years. Each elk that was monitored for more than verse Mercator Coordinate System. For analyses 1 year provided opportunities for an evaluation of and interpretations of summer distributions of the its seasonal range fidelity. elk and their fidelity to specific summer ranges, we We examined whether more than the expected used the same geopolitical herd segment designa­ number of yearlings (sexes combined) summered tion as the state and federal wildlife management in Grand Teton National Park, as reported by agencies (Straley 1968; Cole 1969): Grand Teton Martinka (1969) during the 1960s. We compared National Park, Yellowstone National Park, Teton the proportion of yearlings (sexes combined) of the Wilderness, and Gros Ventre Drainage herd seg­ harvested elk of the Grand Teton National Park ments (Fig. 2). For analyses of migratory patterns, herd segment with the percentage of young (<1- the Grand Teton National Park herd segment was year-old elk) in the censuses during the previous subdivided into a NW Grand Teton National Park winter at the National Elk Refuge. 12 RESOURCE PUBLICATION 199

Data on cattle grazing allotments, dates of per­ studies. These estimates included elk in only a mitted grazing, and number of cattle were ob­ portion of the Grand Teton National Park valley. tained from Grand Teton National Park and the Maximum counts from 2-4 repetitions each sum­ Bridger-Teton National Forest. We mapped sea­ mer were expanded to provide an estimate of the sonal distributions of bison from biotelemetry re­ total number of elk in the valley. The 95% confi­ locations (S. Cain, Grand Teton National Park, dence intervals are approximately ±27% of the unpublished data) and recorded observations by estimates. Boyce (1989) recently provided an personnel of Grand Teton National Park, the Na­ evaluation of the Martinka estimator. He con­ tional Elk Refuge, and the Wyoming Game and cluded that the precision of the estimates seemed Fish Department. reasonable but the results were biased underesti­ mates of the number of elk in the valley because of poor observability of the increasing number of Delineation of Calving Areas adult males. Annual classifications (young, yearling males, Relocations of female elk during the peak par­ mature males, and females) of elk in the Grand turition period of 25 May-15 June 1978-83 Teton National Park central valley were routinely (Johnson 1951; Murie 1951; Oldemeyer et al. conducted with spotting scopes by park personnel 1993) were plotted on U.S. Geological Survey from vantage points in the park. An evaluation of maps (1:24,000 scale) with the Universal Trans­ the reliability of the classification technique is verse Mercator grid system to define areas that forthcoming (S. Cain and B. Smith, in prepara­ female elk frequented during calving seasons. To tion). We also classified elk that we encountered estimate the proportion of the Jackson herd unit during radio-tracking. in which the radio-collared female elk calved, the Annual winter censuses and classifications of number of 1-km2 grid squares in which one or elk on the feed grounds on the National Elk Refuge more females were located during 25 May-15 and in the Gros Ventre drainage were conducted June were divided by the 5,195 km2 of the Jackson in a systematic fashion from feed wagons. After herd unit. Because we did not always see the hay was distributed in two long lines, the feed females that we located by radio and because wagon slowly passed between the two rows of radio-collared females were often with other elk, feeding elk. Young, yearling males, mature males, we could rarely confirm whether a radio-collared and total number of elk were counted by federal­ female had given birth. When a young accompa­ and state-agency biologists and managers. The nied a lone radio-collared female, we assumed the number of females was obtained by subtracting young was her offspring. the number of elk in all other age classes from the We investigated the fidelity to calving areas of total number of counted elk. Replicate counts of females that we monitored during two or more each age and sex class by the multiple observers calving seasons. To do so required delineating were averaged. These counts were scheduled to specific calving areas from relocations of elk. Our coincide with the greatest attendance of the feed definition of calving areas as "the areas, usually grounds by elk (Boyce 1989). on spring range, where females give birth to young and maintain them during their first few days or weeks" follows Thomas et al. (1979:471). Evaluation of Harvests

Harvests of elk of the four herd segments was Estimated Sizes and Age-Class deduced from harvests of radio-collared elk. Har­ Compositions of Herd Segments vest rates of elk of the Jackson herd unit were determined from radio-collared elk and also from Estimates of the number of elk summering in harvest records compiled by the Wyoming Game Grand Teton National Park from 1963 to 1985 and Fish Department, Grand Teton National were made with a Lincoln-Petersen estimator de­ Park, and the National Elk Refuge. These records veloped by Martinka (1969) from mark-recapture are based on (1) an annual mail survey oflicensed MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 13

elk hunters conducted by the University of Wyo­ where p = the proportion ofradio-collared elk from ming under contract to the Wyoming Game and a herd segment and n = the total number of Fish Department, (2) a permanent hunter check radio-collared elk. Because the radio-collared elk station east of the Jackson herd unit along State were assumed to be a random sample of the herd, Highway 26/287 at Dubois, Wyoming, operated the SEs were applied to estimate the number of throughout the elk hunting season, and (3) man­ elk in herd segments. datory reporting of elk killed by hunters that re­ All point estimates of parameters are presented ceive permits to hunt in Grand Teton National ± SE unless otherwise indicated. The levelP <0.05 Park and the National Elk Refuge. Hunters in the was considered significant. park and on the refuge are either contacted by enforcement personnel in the field or must return a portion of their permit with information about Results the results of their hunt.

Statistical Treatment Summer Distribution

Standard statistical procedures described by We monitored the 97 radio-collared elk for an Sokal and Rohlf(1981) were used. Correlation and average of 19.2 months (Appendix A). However, 9 regression analyses were used to test relations premature radio failures and 3 natural mortalities between continuous random variables. Stepwise precluded determining the summer ranges of 12 multiple regressions to predict fidelity to calving of the 97 elk. The remaining 85 animals used all areas were conducted with the following inde­ four summer ranges: 48.2% (95% CI= 37 .6 to 58.8) pendent variables: distance of calving area from summered in Grand Teton National Park, 28.2% the National Elk Refuge, elevation of calving area, (18.6 to 37.8) in Yellowstone National Park, 11.8% and climatic data of calving areas near one of the (4.9 to 18.6) in the Teton Wilderness, and 11.8% four weather-reporting stations in the Jackson (4.9 to 18.6) in the Gros Ventre drainage (Table 3). 2 herd unit. Differences in means were tested with The number of males (X = 8.2, 2 df, P = 0.02), 2 the Student's t test and one-way analyses of vari­ number offemales (X = 25.1, 2 df, P <0.001), and ance. Differences in frequencies and proportions number of the combined sexes of the radio-collared were examined with chi-square and Z-test analy­ elk in each of the four herd segments were not the 2 ses. Because of the relatively small samples of same (X = 30.9; 2 df; P <0.001). monitored, radio-collared elk in any given year in Of the 85 monitored elk, 45.8% ±10.6 (95% CI) any given herd segment of either gender, prob­ summered in areas that were closed to hunting abilities of Type II errors were high in hypothesis west of the Snake River and Jackson Lake in tests. When appropriate, data from years or herd Grand Teton National Park (Figs. 2, 3). Only 2.4% segments were sometimes combined. of the 85 elk summered in areas that were open to Maximum-likelihood survival rates of adult hunting (Hunting Units 75, 76, and 79; Fig. 3) in (~2.5 year-old) elk were estimated from the bio­ Grand Teton National Park. telemetry data with the Program SURVIV (White 1983). Because elk were captured and radio-col­ lared in March or April, annual survival rates Fidelity to Winter Ranges were measured from 1 April of the year in which the animals were collared to 1 April of the succeed­ The fidelity to the National Elk Refuge winter ing years. range was 97% (138 of 142 opportunities) among Standard errors (SE) of the proportions of ra­ the radio-collared elk that survived one or more dio-collared elk in each herd segment were calcu­ years. Three of the four times when elk did not lated with the estimated variance for the binomial return to the National Elk Refuge (but remained distribution: in the Jackson herd unit) occurred during win­ SE= -vp (l-p)ln ter of 1980-81, when snowfall was so low that 14 RESOURCE PUBLICATION 199

Table 3. Distributions of 85 radio-collared elk (Cervus elaphus) of the Jackson elk herd and the number of summers during which the elk were monitored, 1978-1984. Data are presented by herd segment, each of which is identified by its summer range. Number of monitored elk Number of summers of monitoring Herd segment Total ---1 2 3 4 5 (summer range) Male Female n % M F M F M F M F M F Northwest Grand Teton 2 4 6 7.1 1 2 1 2 0 0 0 0 0 0 National Park Grand Teton National 7 28 35 41.2 1 11 5 3 0 4 1 5 0 5 Park valley Combined a 9 32 41 48.2 2 13 6 5 0 4 1 5 0 5 Yellowstone National Park 3 21 24 28.2 2 11 1 6 0 2 0 0 0 2 Teton Wilderness 4 6 10 11.8 3 2 1 0 0 4 0 0 0 0 Gros Ventre drainage 1 9 10 11.8 0 4 1 3 0 1 0 1 0 0 Total 17 68 85 100.0 7 30 9 14 0 11 1 6 0 7 a Northwest Grand Teton National Park and Grand Teton National Park valley.

supplemental feeding was not required at the Na­ 2.3, n = 20) in Grand Teton National Park during tional Elk Refuge. summer (Table 4) and have been significantly higher than yearling-male-to-100-female ratios on the National Elk Refuge during subsequent win­ Fidelity to Summer Ranges ters ( x = 10.l ±0.7, n = 17, t = 4.83, P <0.001). Furthermore, an average of 86 ±2.4 (n = 18) elk Radio-Collared Adult Elk failed to migrate each spring and remained on the National Elk Refuge in summer. In some years, The fidelity to summer ranges was 98% by 48 more than 50% of them were yearling males; 66% elk that were monitored during 2-5 summers. The (177 of 270) of elk harvested during the first week two exceptions were a female that summered in of the National Elk Refuge hunting seasons since Yellowstone National Park in 1980 and 1981 but 1978 have been yearling males. New young were in Grand Teton National Park in 1982 and a fe­ not seen on the refuge in summer, and many male that summered in the Teton Wilderness in antlerless elk were yearling females. 1978 and 1979 but in the Gros Ventre drainage in We have no data to clarify whether any of these 1980. yearlings migrated as 2-year-olds to summer The fidelity of most animals went beyond re­ ranges outside Grand Teton National Park. A turning to one of the four general summer ranges. greater proportion of 3-year-old females than For example, 10 of the 11 radio-collared elk (moni­ either yearling females (F = 4.4; 1, 19 df; P = 0.05) tored ;;::2 years) that summered in Yellowstone or 2-year-old females (F = 7.7; 1, 18 df; P = 0.002) National Park returned to the same drainage or have been harvested in Grand Teton National mountain complex each summer. Park since 1977. Yearlings The percentage of yearlings (sexes combined) Spring Migration among the annual harvests of elk from Grand Teton National Park was significantly higher (F = 4.4; 1, 18 df; P = 0.05 since 1977; F = 3.2; 1, 34 df; Departures of radio-collared elk from the Na­ P = 0.08 since 1969) than the percentage of young tional Elk Refuge in spring peaked approxi­ during classifications at the National Elk Refuge mately 3 weeks after the termination of supple­ in previous winters. Since 1969, yearling-male-to- mental feeding, which depended on snowmelt and 100-female ratios have averaged 21.6 to 100 (SE= consequent greening of forage. Males departed MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 15

Table 4. Age- and sex-class ratios of elk (Cervus elaphus) of the Jackson elk herd on the National Elk Refugea, Wyoming, and at the three feed grounds in the Gros Ventre drainage\ Wyoming, in winter (February) and in the central valley of Grand Teton National Parke in summer. All counted elk on the National Elk Refuge and at the Gros Ventre drainage feed grounds were classified by age and sex.d Summer classificationsd Winter classificationsd Grand Teton National Park valley National Elk Refuge Gros Ventre Estimated MM:100 YM:100 Y:100 Total MM:100 Total MM:100 YM:100 Counted Classified total Year F F F elke F elke F F Y:F elk elkf elkg 1969 16.0 10.4 40.2 9,205 2.8 3,224 21 30 30 794 372 1,431 1970 15.9 9.8 29.4 8,421 2.8 2,334 31 15 32 1,315 586 1,733 1971 14.2 11.2 30.0 8,054 3.3 2,816 41 24 31 1,037 243 1,131 1972 15.1 11.1 33.1 7,615 2.5 2,362 48 19 22 765 331 1,611 1973 15.6 8.8 27.2 7,194 3.8 1,617 14 9 36 899 393 1,644 1974 13.7 10.6 26.2 7,878 1.9 1,729 64 36 16 1,382 279 1,991 1975 15.6 10.9 29.7 7,450 3.0 1,366 43 18 38 1,323 615 1,762 1976 20.7 10.8 34.9 7,858 4.6 l,41~ 102 18 29 947 399 1,076 1977h 17.5 13.1 32.6 5,732 NCO' 165 21 37 1,316 537 1,442 1978 27.5 7.4 30.0 8,413 6.1 1,703 172 38 47 2,718 636 1,800 1979 34.6 12.5 33.0 7,828 2.6 1,686 104 18 36 1,512 934 1,667 1980 37.8 9.9 26.7 7,749 5.1 2,009 200 29 33 1,347 449 1,558 1981h NCOi NCoi 182 20 53 1,744 320 1,396 1982 33.2 10.7 28.0 6,530 2.9 2,095 171 42 49 l,70~ 398 1,753 1983 33.8 14.7 29.0 5,878 0.8 1,442 NC01 1984 37.2 12.0 24.5 5,010 2.2 1,574 310 10 69 1,153 240 1,453 1985 28.0 8.4 28.0 5,758 1.4 1,328 85 15 39 1,331 599 1,804 1986 20.3 8.7 30.3 6,430 1.5 1,665 154 17 52 791 791 836k 1987 19.0 11.6 29.3 7,820 1.5 1,227 54 19 50 1,138 558 1,362 1988 19.3 12.0 30.8 7,753 2.0 1,567 114 21 71 1,122 336 1,180 1989 21.0 11.0 34.0 9,486 2.0 2,334 122 18 55 1,871 760 1,909 Mean 22.8 11.3 30.4 7,403 3.2 1,868 110 22 40 1,192 489 1,527 a Data from this study. bWyoming Game and Fish Department, unpublished job completion report. c R. P. Wood, unpublished data. dF =females> 1-year-old; MM= mature male; YM = yearling male; Y = <1-year-old young of either sex. e The number of elk that were classified on feed grounds. f The number of the counted elk that were classified. gThe number of elk that summered in the central valley of Grand Teton National Park based on a Lincoln-Peterson estimator (Martinka 1969). ~Elk were not supplementally fed during these winters because oflimited snow accumulations. 1 NCO = No classification obtained. j No count because of adverse weather. k A low count was recorded because of fog and poor visibility. 16 RESOURCE PUBLICATION 199

Most elk of the Jackson elk herd migrate from the National Elk Refuge, Wyoming, to their summer ranges in April and May; May 1984.

Elk on the summer range in the central valley of the Grand Teton National Park; July 1985. Mount Moran in the background. MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 17

Males

1.00 ------1-~~ ---;;--;2-­ !'_March ~ April D May ~ Ju_rie

(I) CJ) n = 11 ::, 0.75 ~ n = 17 CJ) -~C: ~ 0.50- c: n = 30 :e0 0 0.25 a.e

Fig. 4. Spring migration of radio-collared 1980 1981 1982 1983 1980-83 elk from the National Elk Refuge, Year 1980-1983. Supplemental feeding ended 1 April 1980, 16 April 1982, and Females 22 April 1983 and was not necessary ------· ---·------·----~~- in 1981. 1.00 -~~ m;;~OMay ~June I n = 15 - ---1 (I) CJ) n = 31 n = 25 ::, 0.75 ~ n = 25 n = CJ) 96 -~C: ~ 0.50- c: :e0 0 e 0.25 a.

0.00 1980 1981 1982 1983 1980-83 Year significantly earlier than females (X2 = 15.0, 3 df, Fall Migration P <0.005; Fig. 4). Elk departed earliest in spring of 1981 when 40% left in March after a winter with The fall migrations commenced in October or low snowfall and no supplemental feeding. November, and 94% ofthe radio-collared elk arrived Migrations to summer ranges required only a at the National Elk Refuge by 15 December (Fig. 5). few days by animals that summered on nearby The durations of fall migrations varied with dis­ ranges in Grand Teton National Park or the Gros tances between the summer ranges and the Na­ Ventre drainage. Most female elk that summered tional Elk Refuge (Table 5). The Grand Teton Na­ in the Teton Wilderness and Yellowstone National tional Park valley and the Gros Ventre drainage Park reached their summer ranges in late June herd segments spent significantly less time (F = and July. The migrations of most females were 13.8; 4, 89 df; P <0.01; x = 8.2 days) in migration interrupted by calving. Male elk arrived at the than the NW Grand Teton National Park, Teton summer ranges in the Teton Wilderness and Yel­ Wilderness, and Yellowstone National Park herd lowstone National Park 3-4 weeks earlier than segments ( x= 18.5 days). Some elk that summered female elk. in southern Grand Teton National Park and the 18 RESOURCE PUBLICATION 199

1983 1982 1981 YNP 1980 1979 1978

1982 1981 1980 TW 1979 1978

a: 1982 1981 LiS 1980 GROS 1979 VENTRE >- 1978

1982 1981 NW 1980 GTNP 1979 1978

1983 1982 VALLEY 1981 GTNP 1980 1979 1978 I OCT24 OCT31 NOl/7 NOV15 NOV22 NOV30 DEC7 DEC15 DATE Fig. 5. Mean dates of departure (left ends of solid bars) of radio-collared elk from summer ranges in Yellowstone National Park (YNP), Teton Wilderness (TW), Gros Ventre drainage (GV), Northwest Grand Teton National Park (NW GTNP), and Grand Teton National Park (GTNP) valley, and mean dates of arrival (right ends of solid bars) at the National Elk Refuge, 1978-83. Lengths of solid bars represent the mean annual duration of migration. western Gros Ventre drainage migrated to the Na­ centrated elk in places. If heavy snows preceded tional Elk Refuge in 1 day or less time. Elk that or coincided with migration, elk took more di­ summered in the eastern Gros Ventre drainage, rect routes and tended to follow drainages and Teton Wilderness, NW Grand Teton National Park ridgelines. (the area west of Jackson Lake), and Yellowstone Migration routes of 5 elk of the Teton Wilderness National Park took from several days to 8 weeks to herd segment (1 male and 4 females) and 15 elk of reach the National Elk Refuge after leaving their the Yellowstone National Park herd segment summer ranges. (2 males and 13 females) crossed Grand Teton Na­ tional Park. Five female elk of the Teton Wilderness Migration Routes herd segment and nine female elk of the Yellow­ During fall migrations, the elk funneled south­ stone National Park herd segment migrated east of ward from their expansive summer ranges to the Grand Teton National Park. After crossing the Buf­ National Elk Refuge (Fig. 6). Topographic features falo Fork River, some animals remained east of such as Jackson Lake and steep escarpments con- Grand Teton National Park until they reached the MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 19

Table 5. Mean duration of fall migration ofradio-collared female elk (Cervus elaphus) by herd segment of the Jackson elk herd, 1978-1984. The mean duration of migrations was compared between herd segments with one-way ANO VA and least-significant-difference tests. a Herd segment (summer range) Migration duration Yellowstone Gros Northwest Grand Teton Herd segment (days) National Teton Ventre Grand Teton National Park (summer range) iJb x SE Park Wilderness drainage National Park valley Yellowstone National Park 28 20.61 2.18 * * * Teton Wilderness 9 25.22 3.88 * * * Gros Ventre drainage 9 9.33 1.89 NSD Northwest Grand Teton National Park 4 12.25 4.46 * Grand Teton National Parkvalley 47 7.96 1.08

a Asterisk(*) indicates test results were significant at P < 0.01; NSD indicates no significant difference. b Includes all migrations of elk that were monitored for more than 1 year.

National Elk Refuge, whereas others crossed Grand dates of elk from the other herd segments did not Teton National Park in Hunting Unit 76 (Fig. 3). differ by sex (Z = 1.13, P = 0.13; Table 7). Arrival at the National Elk Refuge Timing of Fall Migrations The mean annual arrival dates ofradio-collared The mean annual departure dates of radio-col­ elk highly correlated with the arrival of the entire lared elk from their summer ranges during 1978- Jackson elk herd at the National Elk Refuge 83 inversely correlated with October precipitation (r = 0.946, 6 df, P <0.01; Fig. 7). Radio-collared elk at Moose, Wyoming (r = -0. 79, 5 df, P = 0.06). that summered in Grand Teton National Park Including mean October temperatures as a second arrived significantly earlier (i.e., 6-15 days; F = independent variable did not increase the predict­ 16.9; 3, 137 df; P <0.005) at the National Elk ability of departure dates (r = 0.77, 5 df, P = 0.22). Refuge than elk from the other summer ranges Although the elk departed from their summer (Table 6). Radio-collared elk from the Grand Teton ranges earlier in years when precipitation was National Park valley (all areas except northwest­ greater, they did not arrive significantly earlier at ern Grand Teton National Park) arrived earlier the National Elk Refuge (r = -0.69, 5 df, P = 0.13). (t = 2.31, 86 df,P<0.05) at the National Elk Refuge However, the later elk departed from the more than elk that summered in northwestern Grand distant summer ranges in Yellowstone National Teton National Park. Park, in the Teton Wilderness, and in northwestern A significantly greater proportion of male (0.72) Grand Teton National Park, the fewer days they and female (0.49) elk that summered in Grand Teton National Park than males (0.29) and fe­ spent migrating to the National Elk Refuge (r = males (0.13) that summered outside Grand Teton 0.41, 5 df, P < 0.01; Fig. 5). National Park arrived at the National Elk Refuge The arrival of radio-collared females from the before 23 November (males: Z = 2.00, P = 0.023; Teton Wilderness and Yellowstone National Park females: Z = 4.07, P <0.001). Through 30 Novem­ herd segments at the refuge inversely correlated ber, 80% of the elk that arrived at the National Elk with snow depths on 30 November at Moose, Wyo­ Refuge were from Grand Teton National Park ming (r = -0. 762, 6 df, P <0.05), and at Moran, (Table 7). Wyoming, (r = -0.832, 6 df, P <0.05). Correlations Of the Grand Teton National Park herd segment, increased when radio-collared males were in­ males arrived earlier at the National Elk Refuge cluded in the analyses (Moose: r = -0.949, 6 df, P than females (Z = 1.79, P = 0.04). The mean arrival <0.01; Moran: r = -0.946, 6 df, P <0.01). Likewise, 20 RESOURCE PUBLICATION 199

Yellowstone Lake LEGEND Highways

Feed ground Migration * Routes -·-- VVlnter Range

Lakes

0 10 ... .. '

• J Km \

Fig. 6. Migration routes in the Jackson herd unit (adapted from Boyce 1989).

the mean arrival dates of all elk at the refuge tional Elk Refuge did not correlate with snow (based on dates when 50% of the wintering popu­ depths at Moose (Grand Teton National Park: r = lation reached the National Elk Refuge) correlated -0.412, 6 df, P >0.10; Gros Ventre drainage: r = with snow depths on 30 November during 1969-85 0.299, 6 df, P >0.10) and Moran (Grand Teton (Moose: r = -0.790, 16 df, P <0.01; Moran: r = National Park: r = -0.680, 6 df, P > 0.05; Gros -0.786, 16 df, P <0.01). The arrivals of radio-col­ Ventre drainage: r = 0.298, 6 df, P >0.10). Snowfall lared elk of the Grand Teton National Park and accumulations and annual precipitation increase Gros Ventre Drainage herd segments at the Na- from south to north in the study area (i.e., from MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 21

--- ··---·-·· JAN 6 -- £NIIRE WINIER HlRD -··•---- YNP & 1W RADIOED EU DEC 30 - . --e- .

GI NP RAIJIOFIJ [LI( DEC 23 - - --o- - ALL RAD\Olll ELI< w [!, IJEC 16 -- ~ SNOW IJEPIH 0 -···----··-

IJEC 9 - c;,- .• '1 , \ ' ,G ... ' ' , \ ' llfC 2 - ~·.," ~ - - -0

NOV 25 -

NOV 18-

NOV 11 ...,_ ,,i., -"\. I ·. \ \ .. ' .,...... \ / ·- .,. \ ., .,.• - 40 IL... I . \ / '·· ,, ·- ...,. \ / ··, ... ;I ·,.... - 60

r,q 70 71 72 7.S 74 75 76 77 7R Jq RO Rt R? R3 84 R5 86 YEAR Fig. 7. Relation of30-November snow depths at Moran, Wyoming, to mean arrival dates ofradio-collared elk of the Grand Teton National Park (GTNP), Yellowstone National Park (YNP) and Teton Wilderness (TW) herd segments and all radio-collared elk (Cervus elaphus ), and mean arrival dates of 50% of the total number of elk that wintered at the National Elk Refuge, 1969-1986.

Table 6. Mean 1978-1984 arrival dates at the National Elk Refuge, Wyoming, of radio-collared female elk (Cervus elaphus) of the Jackson elk herd returning from summer ranges. Data are presented by herd segment, each of which is identified by its summer range. One-way ANOVA and pairwise least-significant-difference tests were used to compare dates between herd segments.a Summer range Herd segment Mean arrival dates Yellowstone Teton Gros Ventre Grand Teton (summer range) ~ Calendar Julian (SE) National Park Wilderness drainage National Park Yellowstone National Park 33 5Dec. 339.2 (9.41) * Teton Wilderness 10 6Dec. 340.4 (7.51) * Gros Ventre drainage 12 27 Nov. 331.0 (13.11) * Grand Teton National Park 88 21 Nov. 325.1 (11.78) a Asterisk (*) indicates test results were significant at P < 0.05; NSD indicates no significant difference. b Includes all migrations of elk that were monitored for more than 1 year. 22 RESOURCE PUBLICATION 199

Table 7. Dates of arrival of radio-collared elk (Cervus elaphus) on the National Elk Refuge, Wyoming, on return from summer ranges, 1978-1983. Data are presented by herd segment, each of which is identified by its summer range.

Number(%) of elk a arriving before Herd segment November December January (summer range) Sex ff' 1 16 23 1 16 1 Yellowstone National Park female 30 0 1 3(10) 9(30) 24(80) 29(97) male 3 0 0 0 1 2 Teton Wilderness female 9 0 0 1(11) 2(22) 8(89) 9(100) male 2 le le le le 2 0 Gros Ventre drainage female 9 0 1 2(22) 3(33) 9(100) 0 male 2 0 1 1 2 0 0 Northwest Grand Teton female 6 0 0 0 4 6 0 National Park male 3 0 0 2 3 0 0 Grand Teton National Park valley female 64 1 18 34(53) 52(81) 63(98) 0 male 15 1 7 11(73) 15(100) 0 0 Yellowstone National Park, female 48 0 2 6(12) 14(29) 41(85) 47(98) Teton Wilderness, and male 7 le 2 2 4 7(100) 0 Gros Ventre drainage All Grand Teton National female 70 1 18 34(49) 56(80) 69(99) 69(99) Park male 18 1 7 13(72) 18(100) 0 0 a Data include elk harvested in Hunting Unit 76 of Grand Teton National Park and Hunting Unit 77 of the National Elk Refuge. b Includes all migrations of elk that were monitored for more than 1 year. e This elk was in poor condition and died of natural causes within 2 weeks of its arrival at the National Elk Refuge.

Jackson to Moose to Moran, Wyoming, to the south parture from the summer range and arrival at the entrance of Yellowstone National Park; r = 0.96, df National Elk Refuge) accounted for the later arri­ = 3, P <0.01, and r = 0.99, df = 3, P <0.01). There­ val of elk that summered outside Grand Teton fore, Grand Teton National Park and the Gros National Park (F = 8.88; 1, 12 df; P <0.025). Elk Ventre drainage receive less snowfall during Oc­ that summered in the Grand Teton National Park tober and November than Yellowstone National valley spent much of the time between their de­ Park and the Teton Wilderness. Likewise, there is parture from summer range and arrival at the a gradient in mean annual temperature National Elk Refuge staging on the Baseline Flat from Jackson, Wyoming, to Yellowstone National in Grand Teton National Park (Fig. 1). Prior to Park (r = -0.98, df = 3, P <0.01). Inclusion of crossing the Snake River and completing the mi­ the mean monthly October or mean monthly No­ gration to the National Elk Refuge, hundreds to vember temperatures as a second independent several thousands of elk gathered and foraged variable did not improve the predictability of any each November on sagebrush-grasslands west of models. the Snake River. During fall of 1982 and 1983 when they were consistently relocated every 3 Duration of Migrations days, the elk that summered in the Grand Teton The mean annual dates of departure from the National Park valley spent 77% of their migration summer ranges did not differ between elk that time staging on the Baseline Flat (x migration summered in Grand Teton National Park and elk time = 10. 7 days, x staging time = 8.2 days, that summered on ranges outside Grand Teton n = 26). If their summer ranges included the Base­ National Park (t = 0.115, 10 df, P >0.5) and not line Flat, the elk were excluded from the analysis. between elk that summered in Grand Teton Na­ Some elk of the NW Grand Teton National Park, tional Park valley and elk from all other summer Teton Wilderness, and Yellowstone National Park ranges (t = 0.093, 10 df, P >0.5). Thus, the longer herd segments, which migrated to the National duration of the fall migration (time between de- Elk Refuge through Grand Teton National Park, MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 23

also staged on the Baseline Flat. An insufficient National Park and Teton Wilderness herd seg­ number of radio relocations precluded a compari­ ments used the Pacific Creek and Mt. Randolph son by herd segment of the lengths of time elk calving areas, and only elk of the Gros Ventre spent staging. Drainage segment used the Slate Creek and South Gros Ventre calving areas (Table 8). Radio-collared elk calved in 12 cattle grazing Calving allotments and in the summer range of the Jack­ son bison herd (Figs. 8-10). In five cattle allot­ Calving Locations ments, the grazing season began during the 25 Sixty-five female elk were monitored during May-15 June peak of elk parturition. These in­ 131 calving opportunities (1-4 calving seasons clude the Spread Creek-Uhl Hill cattle allotment each). We collected 449 relocations of those fe­ in Grand Teton National Park, the Fish Creek and males during the peak parturition period (25 Blackrock-Spread Creek allotments in the May-15 June) from 1978 to 1983. The 18 reloca­ Bridger-Teton National Forest, and the Pacific tions not shown on Figure 8 were scattered along Creek allotments in Grand Teton National Park the east side of Jackson Lake, in the Teton Wilder­ and the Bridger-Teton National Forest (Appendix ness, and in southern areas of Grand Teton Na­ D). Although the start of grazing in the Blackrock­ tional Park. Spread Creek allotment was scheduled for 15 During the study, we confirmed calving by 9 June, cattle from the Spread Creak-Uhl Hill allot­ different radio-collared females. Relocations of ment often grazed in this important elk-calving those animals (7 summered in Grand Teton Na­ area earlier in June. tional Park and 2 in Yellowstone National Park) We did not sample characteristics of sites where were typical of the general pattern of relocations we relocated elk. In general, elevations between of all radio-collared fem ale elk from 25 May to 2,000 and 2,500 m, warm aspects (S, SW, or SE) 15 June (Fig. 8). with gentle slopes, and plant communities of sage­ brush-grassland and aspen mixed with coniferous Delineation of Calving Areas forests or-in some areas-willow shrublands The 65 radio-collared female elk frequented were typical of the calving areas. 2 only 8% of the 5,195 1-km grid squares in the Affinities by Herd Segment Jackson herd unit during the peak parturition period. The 65 were a fraction of the total number Ninety-one (69%) of 131 opportunities for calv­ of calving females in the Jackson herd unit. We ing occurred in the Grand Teton National Park delineated seven calving areas from the clumped valley and included elk from every herd segment distribution of relocations (Fig. 8). Most reloca­ (Table 8). In another 17 (13%) calving opportuni­ tions (342) were concentrated in the Grand Teton ties, elk frequented the valley and one other calv­ National Park valley calving area between the ing area. Thus, as many as 82% of the young of Moose townsite and Two Ocean Lake (Fig. 8). In radio-collared elk may have been born in the addition to the calving area in Grand Teton Na­ Grand Teton National Park valley. Sixty-three tional Park valley, 6 other calving areas were (58%) of those 108 calving opportunities were at­ identified at (1) the Snake River in Yellowstone tributable to elk that summered in the Grand National Park, (2) North Jackson Lake, (3) Pacific Teton National Park valley. All radio-collared fe­ Creek, (4) Mt. Randolph, (5) Slate Creek, and (6) males that summered in the Grand Teton National the South Gros Ventre drainage (Fig. 8). Whereas Park valley were relocated in only the Grand Te­ elk from all herd segments used the Grand Teton ton National Park valley calving area during 25 National Park valley calving area, only elk of the May-15 June. Yellowstone National Park herd segment used the Elk from the Yellowstone National Park, Teton Snake River calving area, only elk of the Yellow­ Wilderness, and Gros Ventre Drainage herd seg­ stone National Park and NW Grand Teton Na­ ments used 3-5 of the 7 calving areas (Table 8). tional Park herd segments used the North Jack­ Elk of the Yellowstone National Park herd seg­ son Lake calving area, only elk of the Yellowstone ment used all calving areas that their migra- 24 RESOURCE PUBLICATION 199

...., - Calving area boundaries -- Park boundaries • -• .. National Elk Refuge I Yellowstone 1 0~-~ 2 4 6 8 10 .s\,,f. National Kilometers 6> ,s,/.t-c Park 1 ------,.------_J ...,, /., L < ~ NJL -~.----, 12 Grand Teton National Park I \ J l \ ,...,.J \ J (~ j-"L J

Fig. 8. Number of relocations of radio-collared female elk during the peak parturition period of 25 May-15 June 1978-1983 in each calving area: Snake River (SR), North Jackson Lake (NJL), Pacific Creek (PC), Mount Randolph (MR), Slate Creek (SC), and South Gros Ventre (SGV). The Grand Teton National Park valley calving area is subdivided into six calving centers: Two Ocean Lake (TO), Spread Creek-Uhl Hill (UH), Cow Lake-Signal Mountain (CL), Burned Ridge-Potholes (BR), Mystic Isle Burn (MI), and Taggart Lake-Baseline Flat (TL). MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 25

Table 8. Number of radio-collared elk (Cervus elaphus) of the Jackson elk herd that used the calving areas in the Grand Teton National Park valley, northwest Grand Teton National Park, Yellowstone National Park, Teton Wilderness, and the Gros Ventre drainage from 25 May to 15 June 1978-1983. Data represent the number of radio-collared elk that used each calving area summed across all 6 years and are presented by herd segment, each of which is identified by its summer range. Herd segments (summer range) Grand Teton Northwest Yellowstone National Grand Teton National Teton Gros Calving areas Park valley National Park Park Wilderness Ventre Total Grand Teton National Park 63 3 14 6 5 91 valley Snake River in Yellowstone National Parka 0 0 10 0 0 10 North Jackson Lakea 0 1 4 0 0 5 Pacific Creeka 0 0 5 2 0 7 Mount Randolpha 0 0 3 2 0 5 Slate Creek 0 0 0 0 5 5 South Gros Ventre 0 0 0 0 5 5 Snake River and North Jackson Lake 0 0 1 0 0 1 North Fork Buffalo River 0 0 0 2 0 2 Total 63 4 37 12 15 131 a Some relocations of elk in these calving areas were also made in the Grand Teton National Park valley. No elk used more than 2 areas/year.

tion routes crossed (Valley, Pacific Creek, Mount sequent years and more than the elk of NW Grand Randolph, North Jackson Lake, and Snake River Teton National Park (50%), Teton Wilderness in Yellowstone National Park calving areas), and (43%), and Gros Ventre drainage (0%) herd seg­ elk of the Teton Wilderness used all calving areas ments. Fidelity inversely correlated with distance that their migrations routes crossed (Valley, Pa­ between the winter range and each calving area cific Creek, and Mount Randolph calving areas). (r = -0.83, 6 df, P <0.05). Five of 15 elk of the Gros Ventre drainage More female elk of the Yellowstone National herd segment spent the calving season in the Pot­ Park herd segment used the Snake River calving holes, Signal Mountain, or Spread Creek area (the highest in elevation and farthest from of the Grand Teton National Park valley before the National Elk Refuge) in 1981 than in other migrating southeast to their summer ranges years of the study. During this study, the snowpack (Figs. 2 and 8). melted and vegetation began greening earliest in spring of 1981. Of three elk of the Yellowstone Fidelity to Calving Areas National Park herd segment that spent the 1981 The fidelity to calving areas was 78% (50 of 64 calving period at the Snake River calving area, opportunities) among the radio-collared female only one calved there the previous year. Of five elk elk that were monitored during two or more suc­ of the Yellowstone National Park herd segment cessive calving seasons (Table 9). The fidelity to that were monitored during the 1981 calving sea­ calving areas was 100% by the female elk that son, two returned to the Grand Teton National summered in the Grand Teton National Park val­ Park valley calving area and one returned to the ley but 55% by the female elk that summered Snake River calving area in 1982. The other two elsewhere. The elk of the Yellowstone National female elk spent the 1981 calving season at Park herd segment were least loyal. They the Snake River calving area but used the more changed calving areas 67% of the time in sub- southern Grand Teton National Park valley and 26 RESOURCE PUBLICATION 199

-- Park boundaries '""' • - • - Nat. Elk Refuge I - Summer and Spring/Fall Ranges I of the Jackson bison herd I Yellowstone ~ ~ 1 0246810- National Kilometers Park 1 ______J

Fig. 9. Summer and spring-fall ranges of the Jackson bison herd. MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 27

....., -- Park boundaries • - • - National Elk Refuge Heart I --Cattle allotments Lake -+ -+ Cattle driveways I ~ -: Yellowstone : 0 2 4- 6 8 10 National Kilometers Park I ______J

( - I \ Grand J Teton 1 National \Park ,..,..J \ J (" j"l. J

Fig. 10. Cattle allotments (capital letters) and cattle driveways (arrows) in the Jackson elk herd unit. Appendix D lists grazing dates and stocking rates of each allotment. 28 RESOURCE PUBLICATION 199

Table 9. Fidelity to calving areas (based on the area in which most relocations were made from 25 May to 15 June) by radio-collared female elk (Cervus elaphus) of the Jackson elk herd, Wyoming. Only elk monitored during 2 or more successive calving seasons are included. Calving area in ;year X + 1 Grand North South North Calving area Teton Snake Jackson Pacific Mount Slate Gros Fork inyearX valley River Lake Creek Randolph Creek Ventre Buffalo Total Grand Teton valley 40 2 1 2 2 0 0 0 47 Snake River 1 2 1 0 0 0 0 0 4 North Jackson Lake 1 1 0 0 0 0 0 0 2 Pacific Creek 1 1 0 0 0 0 0 0 2 Mount Randolph 1 0 0 0 1 0 0 0 2 Slate Creek 0 0 0 0 0 3 0 0 3 South Gros Ventre 0 0 0 0 0 0 3 0 3 North Fork Buffalo 0 0 0 0 0 0 0 1 1 Total 44 6 2 2 3 3 3 1 64

North Jackson Lake calving areas in 1982. Spring valley (53%) and elk that summered in all other arrived late in 1982 and supplemental feeding on locations (56%). the National Elk Refuge continued until 16 April Female elk that summered in the Grand Teton (Fig. 4). National Park valley used the southern and west­ Fidelity was best predicted by a multiple re­ ern calving centers most. Female elk from the gression of three independent variables: elevation Yellowstone National Park and Gros Ventre herd of the calving area, distance of the calving segments frequented the northern and eastern area from the National Elk Refuge, and mean calving centers most. Elk that summered in north­ annual temperature (R 2 = 0.99, 4 df, P = 0.04). west Grand Teton National Park, which migrate Mean annual precipitation and snowfall did along the west side of Jackson Lake to reach their not significantly contribute to any regression summer ranges, were found in only the Mystic Isle equations. Burn and Burned Ridge-Potholes calving centers (Table 10). Grand Teton National Park Valley Calving Area Natality The most intensively used calving area was the Grand Teton National Park valley. Relocations Pregnancy rates were 17% of 50 1.5-year-old there were clustered in six calving centers, rang­ 2 female elk and 87% of318 ~2.5-year-old female elk ing from 9 to 16 km (Fig. 8). Distributions of 28 that were trapped on the National Elk Refuge elk that spent the calving season in the Grand during 1976-82. The expected natality was 68 Teton National Park valley were restricted to 1 of young/100 female elk (Figure 11). Reducing this these 6 calving centers during any year. The re­ estimate by 7% to account for losses offetuses from maining 33 (54%) elk travelled between two or brucellosis (Oldemeyer et al. 1993), we expected three centers (based on female elk that were relo­ 63 young/100 female elk at birth. cated at least once each week during the 3-week Since 1970, midwinter young-to-100-female-elk calving period). The percentage of animals that ratios at the National Elk Refuge have averaged used more than one calving center in the valley 29.6 ± 2.9 or about 50% of the expected ratios did not differ (Z = -0.18, P = 0.43) between elk (Table 4). Over half of the difference between that summered in the Grand Teton National Park the expected and observed production was lost by MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 29

Table 10. Use of calving centers in the Grand Teton valley calving area by radio-collared female elk (Cervus elaphus) of the Jackson elk herd, Wyoming, during 25 May-15 June 1978-1983. Telemetric relocations of the elk are presented by herd segment, each of which is identified by its summer range and expressed in relative frequency of use. Herd segment Taggart Lake- Mystic Isle Burned Ridge Cow Lake- Spread Creek- 'I\vo Ocean (summer range) Baseline Flat Burn and Potholes Signal Mountain Uhl Hill Lake Total Grand Teton National Park valley 47 50 67 44 3 9 220 Northwest Grand Teton National Park 0 11 4 0 0 0 15 Yellowstone National Park 0 1 6 24 26 24 81 Teton Wilderness 8 0 1 4 2 0 15 Gros Ventre drainage 0 0 2 9 3 1 15 Total 55 62 80 81 34 34 342

midsummer when the young-to-female-elk ra­ young:100 female elk were harvested during falls tios had declined to about 45:100 throughout the 1978-83 (30.3:100 during 1969-86; Boyce 1989, Jackson herd unit (Table 11). The additional decline Appendix C). Either the mortality of young (not (F = 13.8; 1, 8 df; P = 0.006) in the young-to-female­ attributable to hunter selectivity for young) oc­ elk ratios on the National Elk Refuge from midsum­ curred between midsummer and midwinter or the mers 1978-83 to the following winters could not be midsummer young-to-female-elk classifications explained by hunter selection of young. Only 30.1 are biased upward.

Herd composition and pregnancy Birth rate expected the following (PG) rates observed in winter June

Female 100 -=--:::::::..:: 87 mature (87% PG) - 87 mature elk bear 76 young elk 13 yrlg (17% PG) 13 2-year-olds bear 2 young

Young 15~ 15~ (0%PG) 15 yrlg elk bear O young 15()"'

Total 115 ~~ 115 ~~ elk bear 78 young

Ratios 30 young: 78 young: 115 converts to a birth 100 females rate of 68 young: 100 females

Fig.11. Expected and observed ratios ofyoung(1 year old) elk (Cervus elaphus) of the Jackson elk herd, Wyoming. The winter herd composition since 1970 (n = 15) and the pregnancy rates of yearlings (yrlg) and mature females during winters of1976-78 were collected at the National Elk Refuge, Wyoming. 30 RESOURCE PUBLICATION 199

Table 11. Young-to-female(> 1-year-old) ratios of the herd segments of the Jackson elk (Cervus elaphus) herd, 1978-1983. Herd segments are identified by their summer ranges. Herd segment Year (summer range) 1978 1979 1980 1981 1982 1983 Total Grand Teton National Park Young 259 220 411 890 Females 592 544 838 1,974 Ratio 44:100 40:100 49:100 44.4:100 Teton Wilderness Young 89 27 116 Females 179 82 261 Ratio 50:100 33:100 41.3:100 Yellowstone National Park Young 63 243 47 116 469 Females 124 559 117 310 1,110 Ratio 51:100 43:100 40:100 37:100 43.0:100 Gros Ventre drainagea Young 190 268 458 Females 407 457 864 Ratio 47:100 59:100 53.0:100 a Data from G. Roby, Wyoming Game and Fish Department.

Ratios did not differ by herd segment (F3,10 = 1.11; P = 0.4). Young-to-female ratios that we observed in Grand Teton National Park (x = 44.4:100) did not differ (t =0.018, 8 df, P = 0.9) from ratios recorded by personnel of Grand Teton National Park during 1978-83 (x = 43.6:100; Table 4).

,Harvests National Park herd segment (15.8%) were har­ vested at the lowest rates (Table 12). Harvest rates 2 Harvest Rates of Radio-collared Elk differed (X = 5.2, 1 df, P = 0.02) between the sexes of elk that summered outside (male = 50.0%; fe­ During 1978-84, 35 of the 85 radio-collared elk male= 20.3%) Grand Teton National Park. Har­ were harvested. Because some animals were vest rates did not differ between the sexes of elk monitored for two or more hunting seasons, there inside Grand Teton National Park (X2 = 0.5, 1 df, were 175 opportunities for the 85 elk to be har­ P = 0.47, male= 11.1%; female= 18.0%) and not vested. The mean annual harvest rate was 20.0% between all radio-collared males (25.0%) and fe­ (Table 12). The elk that summered in the national males (19.0%; X2 = 1.1, 1 df, P = 0.29; Table 12; parks were harvested at lower rates (25 of 143 = probably because of the small sample sizes). Har­ 17 .5%) than the elk that summered in the national vest rates of males (X2 = 5.2, 1 df, P = 0.02) but not forest (10 of 32 = 24.3%; Z = -1.76, P = 0.04). Elk of females (X2 = 0.1, 1 df, P = 0.73) were signifi­ from the Gros Ventre herd segment were har­ cantly higher outside Grand Teton National Park, vested at the highest rates, and elk from the where the hunting of either-sex elk was the rule, Grand Teton National Park herd segment were than inside Grand Teton National Park. harvested at the lowest rates (Table 12). Harvests in the Jackson Herd Unit, 1978-1983 Harvest of Radio-collared Elk by Sex The lower harvest rate of radio-collared male Males of the Grand Teton National Park herd elk in Grand Teton National Park than outside segment (11.1%) and females of the Yellowstone Grand Teton National Park occurred during years MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 31

Table 12. Number of radio-collared elk (Cervus elaphus) of the Jackson elk herd that were harvested during 1978-1983. Data are presented by herd segment, each of which is identified by its summer range. Harvests of radio-collared elk in Hunting Units 75, 76, and 79 of Grand Teton National Park and in Hunting Unit 77 on the National Elk Refuge, Wyoming, are shown by week. Harvested in Grand Teton National Park and on the National Elk Refuge Herd segment Harvested Before 11/16- 11/24- After Hunting by sex Number3 Number (%) 11/16 11/23 11/30 11/30 season Grand Teton National Park Male 18 2 (11.1) 1 1 0 0 2 Female 83 15 (18.1) 1 5 6 3 15 Both 101 17 (16.8) 2 6 6 3 17 Yellowstone National Park Male 4 2 (50.0) 0 0 0 1 1 Female 38 6 (15.8) 0 1 0 2 3 Both 42 8 (19.0) 0 1 0 3 4 Teton Wilderness Male 4 2 (50.0) 0 0 0 0 0 Female 11 2 (18.2) 0 0 0 0 0 Both 15 2 (26.7) 0 0 0 0 0 Gros Ventre drainage Male 2 1 (50.0) 0 0 0 0 0 Female 15 5 (33.3) 1 0 1 1 3 Both 17 6 (35.3) 1 0 1 1 3 Total Male 28 7 (25.0) 1 1 0 1 3 Female 147 28 (19.0) 2 6 7 6 21 Both 175 35 (20.0) 3 7 7 7 24 Combinedb Male 10 5 (50.0) 0 0 0 1 1 Female 64 13 (20.3) 1 1 1 3 6 Both 74 18 (24.3) 1 1 1 4 7 3 Elk that were monitored for more than one hunting season were available to hunters each year. The sample sizes in the table equal the sum of all years that each elk was available to hunters. bYellowstone National Park, Teton Wilderness, and Gros Ventre drainage combined. when hunting seasons were designed primarily to Grand Teton National Park herd segment. Antler­ harvest antlerless elk in Grand Teton National less elk were 64% of the harvested elk in those Park. Antlered elk have composed 60-70% of the units during 1981-84 (Table 13). The composition harvest with either-sex permits in Hunting Units of the harvested radio-collared elk, standardized 75-77 and 79 (Wyoming Game and Fish Depart­ for sample sizes, was 62% antlerless elk. ment. 1978-86. Annual Big Game Completion Re­ Outside Grand Teton National Park, hunting ports, unpublished). From 1981 to 1984, when seasons were designed for the harvest of primarily male and female elk were radio-collared, antler­ male elk. From 1978 to 1983, elk hunting with less permits composed 72% of the total permits general licenses was only for antlered elk, except issued for Hunting Units 75-77 (Fig. 3; Appendix during a portion of the season when antlerless elk C). The remaining 28% were issued for elk of also could be killed. In Hunting Units 82 and 83 either sex. The emphasis on antlerless elk was to in the Gros Ventre drainage, antlerless elk could expedite the reduction of the number of elk on the be harvested only with limited quota permits (Ap­ National Elk Refuge, particularly those from the pendix C). 32 RESOURCE PUBLICATION 199

Harvests in Hunting Units 75-77 were directed Whereas 73% of the harvested elk in Hunting at the Grand Teton National Park herd segment, Units 75-77 in 1984-86 were males, only 22% were and the antlered-to-antlerless-elk-harvest ratio males in 1978-83. As a result, classification counts (radio-collared elk and all harvested elk) was lower in Grand Teton National Park revealed a decline in there than in the Yellowstone National Park, Teton the mature-male-to-100-female-elk ratios Wilderness, or Gros Ventre Drainage herd seg­ (186 ±23.5 in 1977-84 to 91 ±23.3 in 1985-87; Table ments, which were primarily harvested in Hunting 4; F = 5.28; 1, 7 df; P = 0.055). During the corre­ Units 70, 71, 80, 81, 82, and 83 (Table 13; Fig. 3). As sponding winters, mature-male-to-100-female-elk a result, summer mature-male-to-female-elk ratios ratios on the National Elk Refuge declined from in the Grand Teton National Park central valley 34.0 ±1.5 to 22.4 ±2.8 (F = 16.29; 1, 7 df; P = 0.005). increased from a mean of 46:100 (SE = 9.2) during Throughout 1978-86, male-to-female-elk ratios on 1969-76 (when either-sex-elk permits were the the National Elk Refuge correlated (r = 0.87, 6 df, rule) to a mean of 186:100 (SE= 23.5) during 1977- P <0.01) with the percent of antlerless elk in the 84 (P <0.01; Table 4). harvest from Hunting Units 75-77, which corre­ Post-1975 male-to-female-elk ratios on the Na­ lated (r = 0.96, 7 df, P < 0.01) with the percent of tional Elk Refuge, where the elk of the Grand Teton antlerless permits. National Park herd segment winter, confirm the Location of Harvest by Herd Segment upward trend but not the magnitude in Grand Teton National Park valley male-to-female-elk ra­ During 1979-83, most elk migrated to the Na­ tios. In contrast, mature male-to-female-elk ratios tional Elk Refuge before the close of the hunting on the Gros Ventre feed grounds (where elk from seasons (Table 7; Fig. 7). Of the 17 radio-collared only the Gros Ventre drainage, Yellowstone Na­ elk harvested in Hunting Units 75, 76, and 79 tional Park, and Teton Wilderness herd segments (Grand Teton National Park) and 77 (National Elk winter) averaged only 3:100 (SE = 0.4) and re­ Refuge) prior to 1 December, 14 were from the mained relatively stable (Table 4). Grand Teton National Park herd segment (Table 12). This mirrored the earlier arrival at the Na­ Harvests in the Jackson Herd Unit After 1983 tional Elk Refuge of the Grand Teton National Park herd segment than the elk from other herd seg­ After 1983, hunting regulations in Grand Teton ments (Table 7). All harvested radio-collared elk of National Park (Hunting Units 75, 76, and 79) and the Grand Teton National Park herd segment were on the National Elk Refuge (Hunting Unit 77) were killed in the park or on the refuge. Seven of 18 elk drastically changed (Appendix C). From as many as harvested from the other three herd segments were 8,413 elk during winter 1977-78, the number of elk killed in the park or on the refuge; four of those were on the National Elk Refuge declined to 5,010 during harvested after 30 November (Table 12). winter 1983-84, which was well below the targeted Harvest rates were not different of elk of the population size of 7,500 (Table 4). To increase the Teton Wilderness and Yellowstone National Park number of elk on the refuge, all hunting permits for herd segments that migrated along eastern (na­ Hunting Units 75-77 during 1984-86 were for elk tional forest lands) migration routes than of elk of either sex (antlerless-only-elk permits were migrating along western (Grand Teton National eliminated), and the number of permits was re­ Park) routes (Z = 0.92, P = 0.18). Whereas 7 (25%) duced (Appendix C). Hunting Unit 79 permits were of the 28 radio-collared elk that migrated along reduced from an average of 2,583 during 1978-83 western migration routes were harvested, only 2 to 1,800 in 1984, 1,000 in 1985, and 1,500 in 1986. (13%) of the 15 radio-collared elk that migrated east Antlerless animals were hunted only under lim­ of Grand Teton National Park were harvested. ited-quota permits in Hunting Units 70, 71, 80, 81, 82, and 83 (Appendix C; Fig. 3). These conservative regulations increased the National Elk Refuge Natural Mortality population to 7,820 elk during winter 1986-87 and to 9,486 elk (15,000 total in the Jackson herd unit) During the 6 years we monitored radio-collared during winter 1988-89. elk, 10 of the 97 animals (3 of 17 males and 7 of 80 MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 33

Table 13. Harvests of elk (Cervus elaphus) in the Jackson herd unita and Grand Teton National Park and on the National Elk Refuge, Wyoming, 1950-1986. Harvest in Grand Teton National Park and in the National Elk Refuge Total harvest in Jackson herd unit Hunting units Hunting units 75-77 Antlered Antlerless Unknown 77b 75 76 79c Antlered Antlerless Year Total no. % no. % no. no. no. Total no. no. % 1950 3,638 1,256 42 1,762 58 620 1951 4,810 1,469 36 2,567 64 774 184 1952 2,108 771 42 1,080 58 257 210 27 1953 2,823 957 45 1,166 55 700 67 112 1954 2,594 1,113 46 1,301 54 180 104 1955 3,207 1,234 40 1,848 60 125 310 1956 3,106 1,028 33 2,078 67 0 325 1957 2,252 713 38 1,164 62 375 160 1958 3,057 1,250 41 1,807 59 0 110 1959 1,542 703 46 832 54 7 0 1960 2,074 767 44 977 56 330 0 1961 2,639 2,639 278 1962 1,320 684 55 554 45 82 280 1963 2,996 1,608 56 1,284 44 104 95 625 496 1964 2,975 1,534 53 1,373 47 68 50 753 611 1965 3,250 1,775 56 1,397 44 78 96 691 570 1966 3,226 1,907 59 1,319 41 0 133 612 306 1967 2,974 1,879 63 1,095 37 0 131 375 286 1968 3,090 1,776 43 2,383 57 26 68 520 351 1969 3,444 1,873 54 1,571 46 0 148 568 529 1970 5,680 2,618 46 3,055 54 7 162 828 592 1971 4,245 1,608 38 2,583 62 54 70 584 366 98 268 73 1972 4,086 1,666 41 2,420 59 0 59 519 553 235 318 58 1973 4,684 1,908 41 2,776 59 0 77 571 306 76 230 75 1974 2,774 1,033 38 1,681 62 0 49 451 185 77 108 58 1975 3,526 1,631 46 1,895 54 0 139 701 651 338 313 48 1976 1,429 670 47 759 53 0 0 293 113 0 113 100 1977 3,756 1,799 48 1,957 52 0 220 642 505 0 505 100 1978 2,880 1,131 39 1,749 61 0 227 659 512 0 512 100 1979 3,321 1,225 37 2,096 63 0 280 691 801 34 767 96 1980 3,740 1,847 49 1,893 51 0 513 492 629 312 317 50 1981 4,290 1,964 46 2,326 54 0 481 501 614 163 451 73 1982 3,548 1,343 38 2,205 62 0 323 870 636 110 526 83 1983 2,355 1,165 49 1,190 51 0 317 618 488 124 364 75 1984 1,524 1,087 71 437 29 0 132 372 250 189 61 24 1985 1,368 1,009 56 796 44 0 178 343 335 237 98 29 1986 1,254 977 76 317 24 0 140 344 203 149 54 27 Mean 3,016 1,361 46 1,602 54 173 174 456 454 165 313 67 a5,195 km2 in the Snake River watershed. bThere was not a hunt in Hunting Unit 77 in 1950, 1951, and 1954-62. c Includes harvests from Hunting Unit 72 in Grand Teton National Park 1951-62. Before 1963 Hunting Units 75 and 76 in Grand Teton National Park were not open to hunting. Unit 75 was closed to hunting 1974-76.

females) died of causes believed to be unrelated to 8 of the 10 dead elk were known. Four summered hunting. Six of 10 (2 males and 4 females) died on in Grand Teton National Park and 4 summered in the National Elk Refuge between November and the Teton Wilderness. May. The other four died on summer ranges be­ Causes of death could not be determined from the tween July and November. The summer ranges of skeletal remains. One female elk that summered in 34 RESOURCE PUBLICATION 199

the Teton Wilderness and died on the refuge in April survival between radio-collared elk that summered 1983 had been immobilized during the previous in national parks and those that summered in the month to replace her radio collar. At that time she national forest approached statistical significance was emaciated and suffered from a swollen right­ (X2 = 26.40, 17 df, P = 0.07). Survival did not differ front knee joint and laminitis of a swollen left-rear between males (x = 0.630, n = 18, X2 = 2.503, 1 df, foot on which the hoof was 40 cm overgrown. Judg­ P = 0.11) and females (x = 0.778, n = 69; Table 14). ing by tooth wear, she was > 15 years old. A male that summered in Grand Teton National Park had severe alopecia because of scabies mites (Psorptes Discussion equi cervinus) and was emaciated when he died on the refuge in March 1983. His age from tooth ce­ mentum analysis was 7 .5 years at that time. He did not have clinical scabies when radio-collared in Distribution March 1981. Another male elk that summered in the Teton Wilderness had clinical scabies when he Summer Distribution of Elk died on the National Elk Refuge in November 1981 but not when he was radio-collared in the previous Cole (1969) estimated that in 1964 approxi­ spring. According to tooth wear, he was 3.5 years mately 70% of the elk that wintered on the National old in spring 1981. Elk Refuge summered in Yellowstone National Park and Grand Teton National Park. Fifteen to 20 years later, approximately 76% summered in Yel­ Survival lowstone National Park and Grand Teton National Park (based on the distribution of radio-collared Survival of the radio-collared elk from Grand elk). Although the Yellowstone National Park herd Teton National Park was higher than survival ofelk segment seemingly declined (Z = 18.28, P <0.001) from the other three herd segments combined (X2 = from 42% to approximately 28% of the elk wintering 17.651, 9 df, P = 0.04; Table 14). The difference in on the refuge, the Grand Teton National Park herd

Table 14. Survival rates of elk (Cervus elaphus) of the Jackson elk herd, estimated with Program SURVIV (White 1~83). Data are presented by herd segment, each of which is identified by its summer range. Herd segment (summer range) N Sex sa SE 95%CI Allb 85 M&F 0.754 0.034 0.688-0.820 Allb 68 F 0.778 0.035 0.709-0.847 Allb 17 M 0.630 0.095 0.443-0.816 Grand Teton 32 F 0.803 0.091 0.625-0.981 National Park Yellowstone 21 F 0.819 0.087 0.649-0.989 National Park Teton Wilderness 6 F 0.709 0.158 0.399-1.000 Gros Ventre drainage 9 F 0.654 0.267 0.131-1.000 · Grand Teton and 53 F 0.811 0.060 0.693-0.929 Yellowstone national parks Teton Wilderness 15 F 0.681 0.103 0.479-0.883 and Gros Ventre drainage a Survival rates. b All segments. MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 35

segment has increased (Z = 24.36, P <0.001) from Elk-Logging Study, 1970-1985. Montana Depart­ 29% (Cole 1969) to approximately 48% (this study). ment Fish Wildlife and Parks, unpublished), may be a consequence of matrilineal fidelity to ranges Fidelity to the Jackson Herd Unit by young (Murie 1951; Clutton-Brock et al. 1982). Elk of the Yellowstone National Park herd seg­ This underscores the need to manage herd seg­ ment share portions of their summer range with elk ments on distinct summer ranges. from the northern herd of Yellowstone National Present objectives for the Jackson herd specify a Park, the Shoshone River herd of Wyoming, and the posthunting-season population of 11,000 elk dis­ Sand Creek herd of Idaho (Anderson 1958; tributed as follows: 2,400 at the three Gros Ventre Craighead et al. 1972; Rudd et al. 1983; C. Brown drainage feed grounds, 7,500 at the National Elk 1985 Sand Creek elk: population status, movement, Refuge, and about 1,100 elk on winter ranges other and distribution. Idaho Department of Fish and than those feed grounds. Elk from two or more herd Game, unpublished). Only one radio-collared elk segments attend each feed ground and the other left the Jackson herd unit during 1978-84. In 1981- winter ranges (Figs. 2, 6). The size of each herd 82, that female elk wintered 16 km south of the segment on the summer ranges cannot be inferred National Elk Refuge at the Horse Creek feed from the number of elk at feed grounds. Theoreti­ ground. She returned to the National Elk Refuge in cally all 7,500 elk that winter on the National Elk the following two winters. Based on migrations of Refuge could summer, for example, in Grand Teton radio-collared elk (B. Long, M. Hinschberger, G. National Park, and management objectives would Roby, and J. Kimball. 1980. Gros Ventre cooperative be met. If the state and federal agencies seek to elk study: final report 1974-1979. Wyoming Game restore pre-1950 distributions of elk on summer and Fish Commission, unpublished; this study) and ranges, then it is necessary to establish numeric returns of ear tags (Straley 1968; Boyce 1989), the objectives for herd segments on each summer boundaries of the Jackson herd unit, except per­ range. Hunting seasons in each hunting unit should haps for southwestern Yellowstone National Park, then be designed to achieve the desired size and remain unchanged from earlier delineations (An­ composition of each herd segment. derson 1958; Cole 1969). Fidelity to Seasonal Ranges Migration Strong fidelity to seasonal ranges, demon­ strated by the radio-collared elk, also has been Migrations Through Grand Teton National reported by other authors (Anderson 1958; Park Craighead et al. 1972; Houston 1982; Hershey and The distributions of migrating elk have been Leege 1982; Rudd et al. 1983; C. Brown 1985. estimated since 1945 by counting tracks that cross Sand Creek elk: population status, movement, Highway 26/287 between Moose, Wyoming, and and distribution. Idaho Department of Fish and the Togwotee Pass (Fig. 1). Although numbers of Game, unpublished). The fidelity of the elk to elk and proportions of each herd segment that migration routes and summer ranges has major follow various migration routes may have changed implications for the management of the Jackson during the past 4 decades, the geographical loca­ elk herd. The traditional use of seasonal ranges by tion of the routes have remained much the same individual elk and by distinct herd segments, as Anderson (1958) described them. which is also reported of the migratory Sand Cole (1969) reported that 32% of the elk tracks Creek herd (C. Brown, 1985. Sand Creek elk: to winter ranges in 1949, 42% in 1957, and 75% in population status, movement, and distribution. 1964 crossed through Grand Teton National Park. Idaho Department of Fish and Game, unpub­ Tracks of neither the Gros Ventre herd segment lished) and of various elk herds in Montana (L. J. nor of most elk of the Grand Teton National Park Lyon, T. N. Lonner, J.P. Weigand, C. L. Marcum, herd segment that migrate to the National Elk W. D. Edge, J. D. Jones, D. W. McCleery, and L. L. Refuge are counted because of the location of the Hicks. 1985. Coordinating elk and timber manage­ transects. The track counts reflect east to west ment: final report of the Montana Cooperative distributions of elk of the Teton Wilderness and 36 RESOURCE PUBLICATION 199

Yellowstone National Park herd segments that unpublished), and British Columbia (Morgantini migrate through the Jackson herd unit. Recent and Hudson 1988). analyses of the track-count data revealed that 40% Reduced Migration of Elk from Southwestern of the elk in the Jackson herd unit in the 1950s, Yellowstone National Park 60% in the 1960s, 71 % in the 1970s, and 72% in the 1980s migrated through Grand Teton National Cole and Yorgason (G. F. Cole and I. J. Yorgason. Park to reach their winter ranges (Boyce 1989). 1965. Elk migration study, Jackson Hole elk herd, The remainder migrated through national-forest 1965. U.S. National Park Service and Wyoming lands east of Grand Teton National Park (Fig. 6). Game and Fish Commission, unpublished) re­ The migrations of radio-collared elk in our ported that at least 578 elk migrated from south­ study corroborate the findings of increased migra­ western Yellowstone National Park along the tion of elk through Grand Teton National Park shores of Jackson Lake to the National Elk Refuge during the 1970s and 1980s. Seventy-one percent in 1964. Cole (1969) believed that 800 of the elk of the radio-collared elk that summered in Yellow­ that wintered on the National Elk Refuge migrated stone National Park or in the Teton Wilderness to summer ranges in southwestern Yellowstone during this study passed through Grand Teton National Park. Aerial surveys indicated that mi­ National Park. Bendt (1962), Cole (1969), Johnson gration had dwindled to 100-150 elk by the late (1976), and Toman (1987) attributed the decline of 1970s and was no larger during the 1980s (G. Roby, elk that summer in the eastern Teton Wilderness Wyoming Game and Fish Department, and and in eastern Yellowstone National Park to dif­ R. Wood, Grand Teton National Park, personal ferential harvests along the eastern migration communication). No locations ofradio-collared elk routes that increased from improved vehicle ac­ were obtained west of Highway 89/287 in Yellow­ cess for logging in the Bridger-Teton National stone National Park during this study. Forest. Excessive elk harvests have not been reported along migration routes between southwestern Yel­ Behavioral Responses to Hunting lowstone National Park and Grand Teton National Park. However, the apparent decline of migrating Elk of the Grand Teton National Park and Yel­ elk from southwestern Yellowstone National Park lowstone National Park herd segments that sum­ during the 1970s coincided with an increase in the mered adjacent to national-forest lands did not number of wintering elk in the adjacent Sand Creek cross park boundaries until migration began. Fur­ area of eastern Idaho (C. Brown, 1985. Sand Creek thermore, the only two elk that summered in the elk: population status, movement, and distribution. Teton Wilderness near the southern border of Yel­ Idaho Department ofFish and Game, unpublished). lowstone National Park moved into refuge areas Empirical evidence of a shift in migration routes in late summer. One female elk moved 0.5-9.0 km and winter ranges of elk from southwestern Yellow­ north into Yellowstone National Park between 9 stone National Park is lacking. and 22 September during each of 5 years. The other moved 1-8 km into Yellowstone National Implications for Harvests Park between 10 and 22 September during each of As reported of other elk herds (Craighead et al. 3 years. Hunting seasons in the Teton Wilderness 1972; Adams 1982; Hershey and Leege 1982; Rudd opened on 10 September each year, and Teton et al. 1983; C. Brown, 1985. Sand Creek elk: popu­ Wilderness outfitters have noted similar behavior lation status, movement, and distribution. Idaho in elk during mid-September (R. Johnson, outfit­ Department of Fish and Game, unpublished), fall ter, Kelly, Wyoming, personal communication, migrations were stimulated by snow accumulations 1989). Instances of elk moving from areas open to at high elevations on summer ranges. Elk from the hunting to adjacent refuge areas at the onset of more distant summer ranges in Yellowstone Na­ hunting seasons have been reported in Montana tional Park, the Teton Wilderness, and northwest (Knight 1970), Idaho (C. Brown, 1985. Sand Grand Teton National Park arrived significantly Creek elk: population status, movement, and dis­ later (6-15 days) at the National Elk Refuge than tribution. Idaho Department of Fish and Game, elk from the Grand Teton National Park valley MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 37

because of the longer duration of the migration. calving area during 1978-83. Another 13% used This provided an opportunity to harvest primarily the Grand Teton National Park valley and an ad­ elk of the Grand Teton National Park valley herd jacent calving area. This apparent attraction to the segment in Hunting Units 75, 76, and 77 during the Grand Teton National Park valley calving area has early stages of migration. several possible explanations: 1. The expansive, snow-free mosaic of aspens, sagebrush, conifers, and meadows in Grand Te­ Calving ton National Park provides suitable habitat for calving (Skovlin 1982). Similar habitat was less Fidelity to Calving Areas abundant in the other calving areas. The valley The fidelity ofradio-collared elk to calving areas was mostly free of snow by mid-May and pro­ was related to each calving area's elevation and vided ample green forage for pregnant and lac­ distance from the winter range and to annual tating females. Metabolizable energy require­ weather patterns. In years with early snowmelt, as ments of pregnant females increase markedly in 1981, more elk may calve in Yellowstone National during late pregnancy. Requirements continue Park, northern areas of Grand Teton National Park, to rise until about 48 days postpartum when and the Teton Wilderness. The interaction of eleva­ energy expenditure for lactation exceeds main­ tion and temperature probably influenced winter tenance requirements by nearly 60% (Moen snowpack, spring soil temperatures, and conse­ 1973; Nelson and Leege 1982). quent greening of vegetation. Greening is not pre­ 2. The six calving centers in the Grand Teton vented by a lack of soil moisture because moisture National Park valley calving area were in the from snowmelt and precipitation are abundant dur­ summer ranges of the Grand Teton National ing May, the second wettest month of the year. Park valley herd segment (Fig. 8). Ai3 reported by Brazada (1953), residual snow­ 3. Spring migrations of most elk that summered in pack in spring alone did not deter migration of northwest Grand Teton National Park, Yellow­ some elk to calving areas. We observed elk crossing stone National Park, and the Teton Wilderness snow-covered areas during migration to reach ar­ passed through the Grand Teton National Park eas with new vegetative growth. valley. If snows were still deep farther north, Results of our investigation of the Yellowstone which is normal in Jackson Hole, a larger pro­ National Park, Teton Wilderness, and Gros Ventre portion of those elk calved in Grand Teton Na­ Drainage herd segments support Murie's (1951) tional Park valley than farther north. Variation conclusion that most young are born somewhere in calving locations because of late snowmelt between the winter and summer ranges. In re­ and greening of vegetation was previously re­ sponse to the growth in the Grand Teton National ported of elk in Jackson Hole (Altmann 1952; Park herd segment since 1950 and the short migra­ Anderson 1958; and Cole 1969), Montana tion time between their winter and summer ranges, (Knight 1970; Sweeney and Steinhoff 1976), however, many elk now calve on their summer Idaho (C. Brown, 1985. Sand Creek elk: popula­ ranges. Calving on the National Elk Refuge, as tion status, movement, and distribution. Idaho reported by Murie (1951) and Altmann (1952), was Department of Fish and Game, unpublished), not observed during this study and has probably and Yellowstone National Park (Houston 1982). been rare since the 1960s (unpublished National 4. In 1958, grazing of livestock ceased in calving Elk Refuge annual narrative reports). Most elk that areas west of the Snake River in Grand Teton remained on the National Elk Refuge during sum­ National Park. Several authors reported that mer were 1-year-old individuals. elk avoid cattle and that cattle displace elk (Skovlin et al 1968; T. N. Lonner, 1975. Elk-cat­ Grand Teton National Park Valley Calving tle distribution and interspecific relationships. Area Long Tom Creek Study, Montana, Montana Co­ Although we identified seven calving areas dur­ operative Elk/Logging Study Annual Pro­ ing this study, 69% of the radio-collared female elk gress Report, unpublished; Sauer 1980; used only the Grand Teton National Park valley L. J. Lyon, T. N. Lonner, J. P. Weigand, 38 RESOURCE PUBLICATION 199

C. L. Marcum, W. D. Edge, J. D. Jones, D. W. Sympatry and Diseases of Wildlife and McCleery, and L. L. Hicks. 1985. Coordinating Livestock elk and timber management: final report of the Montana Cooperative Elk-Logging Study, 1970- Elk, bison, and domestic cattle are susceptible 1985. Montana Department Fish Wildlife and to brucellosis-the only known, reportable (Code Parks, unpublished). of Federal Regulations, Title 9, Part 161.3(D) dis­ 5. With the exception of early-season tourists along ease of the Jackson elk herd that is potentially roadways and points of interest, Grand Teton transmissible to livestock and humans (Thorne National Park provides elk with security from et al. 1991; Smith and Roffe 1994). The most im­ human disturbance. portant clinical sign of this bacterial infection in elk, bison, and cattle is spontaneous abortion Calving Areas Beyond the Grand Teton (Thorne et al. 1982; Davis et al. 1988). Elk and National Park Valley bison of the Yellowstone ecosystem harbor The Grand Teton National Park valley calving Brucella abortus biotypes 1, 2, and 4-the same area was used by female elk from every summer bacterium and biotypes that are responsible for herd segment, but the other six calving areas were most bovine brucellosis (Thorne et al. 1991). A used exclusively by elk that summered outside the national program to eradicate brucellosis was in­ Grand Teton National Park valley. Consequently, itiated in 1935 by the U.S. Department of Agricul­ preserving the integrity and security of those calv­ ture. The department places commerce restric­ ing areas (B. Long, M. Hinschberger, G. Roby, and tions on livestock operators whose animals J. Kimball. 1980. Gros Ventre cooperative elk contract the disease. Infected livestock herds must study: final report 1974-1979. Wyoming Game be quarantined and retested after 60 days, and and Fish Commission, unpublished; Kuck et al. subsequent reactors must be slaughtered. Ship­ 1985) is paramount to the maintenance of the ment of animals from the herd, except for slaugh­ ter, is prohibited. Yellowstone National Park, the Teton Wilderness, Strain 19, a live vaccine of B. abortus, has been and the Gros Ventre herd segments. The Mount Randolph, Slate Creek, and eastern Spread in use for over 40 years. It is considered 60-70% Creek-Uhl Hill calving areas are of special concern effective in preventing abortion and 55-65% effec­ tive in averting field-strain infections in cattle because most ofthos,e lands do not have the formal that were vaccinated as young (Manthei 1952, protection of national-park and national-wilder­ 1959). Tests and slaughter of reactors and vacci­ ness-area designations. nation have been the only successful means of The North Jackson Lake or Yellowstone Na­ eliminating brucellosis in infected herds (Manthei tional Park Snake River calving areas were not 1959; Davis et al. 1991). reported in previous accounts of the Jackson elk herd (Altmann 1952; Anderson 1958; Murie and Brucellosis in Elk Murie 1966). Elk from winter ranges other than Brucellosis was first diagnosed in 3 of 9 se­ the National Elk Refuge calve in additional areas rologically tested elk in Jackson Hole in 1930 of the Bridger-Teton National Forest and southern (Murie 1951). Since then, free-ranging elk have Yellowstone National Park. North (D. North, 1991. represented a potential source of brucellosis for The Buffalo Valley Elk Enhancement Project. sympatric domestic livestock in late pregnancy or 1990 Annual Report, Wyoming Game and Fish at parturition. The average infection rates since Department, unpublished) captured and radio­ 1970 have been 39% of females and 28% of all collared 11 female elk on their winter ranges 30- wintering elk on the National Elk Refuge (E. T. 35 km north of the National Elk Refuge in the Thorne, unpublished in Boyce 1989). Blood tests Buffalo River valley. During 1990, 9 of the 11 and tissue cultures also revealed brucellosis in elk female elk calved-2 in southeastern Yellowstone or their aborted fetuses at 18 state-operated feed National Park, 1 south of the Teton Wilderness, 4 grounds in western Wyoming (Thorne et al. 1991). in the eastern the Teton Wilderness, and 2 others Only elk in the northern range of Yellowstone probably also in the Teton Wilderness. National Park (1.7% of 6,027 tested elk during MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 39

Intentional or incidental feeding of elk with cattle on private lands during late winter and during spring increases the risk ofinterspecific transmission of brucellosis in western Wyoming; February 1992.

1960-68, M. Meagher, Yellowstone National Park, Wyoming Game and Fish Department, and personal communication, 1989), which are sympa­ Bridger-Teton National Forest. 1994. The Jackson tric with infected bison, maintain the disease in bison herd: long-term management plan and envi­ the absence of winter feeding programs (Smith ronmental assessment. Grand Teton National and Roffe 1994). Park, unpublished). In 1988 and 1989, 76% of 35 A 50% probability exists that a female elk bison that were culled from the herd were serologi­ aborts her first pregnancy after infection with cally positive or suspected of having Brucella an­ brucellosis. Thereafter, she will probably repro­ tibodies. Of the serologic reactors that were cul­ duce successfully but will remain a carrier of the tured for B. abortus, 36% were positive (Williams disease and continue to shedB. abortus in vaginal et al. 1993). The seroprevalence in bison from the exudates, live births, and afterbirths (Thorne northern range of Yellowstone National Park is et al. 1982). This represents a 7% (Oldemeyer 54% (H. I. Pac and K. Frey. 1991. Some population et al. 1993) to 12% (Thorne et al. 1979) reduction characteristics of the northern Yellowstone bison in the annual fecundity. herd during the winter of 1988-1989. Mon­ tana Department of Fish, Wildlife and Parks, un­ Brucellosis in Bison published). There is ample opportunity for inter­ The Jackson bison herd, which numbered 107 specific exposure of bison and elk to B. abortus animals in winter 1987-88, has wintered on the because aborted fetuses of both species have been National Elk Refuge since 1975 and feeds on natu­ observed on the National Elk Refuge (Thorne ral forage and supplemental feed with the elk et al. 1982; Oldemeyer et al. 1993; Williams (Grand Teton National Park, National Elk Refuge, et al. 1993). 40 RESOURCE PUBLICATION 199

Transmission of Brucellosis 1994. The Jackson bison herd: long-term manage­ ment plan and environmental assessment. Grand Transmission of brucellosis from infected elk Teton National Park, unpublished; Smith 1994). and bison to susceptible domestic cows has been Distributions of elk, cattle, and bison overlap on demonstrated (Thorne et al. 1979; Davis et al. many cattle-grazing allotments in Grand Teton 1988). Transmission is possible where infected National Park and in the Bridger-Teton National and susceptible animals are closely associ­ Forest. Cattle also share ranges with calving elk ated.Transmission of brucellosis from infected elk (that winter on the three Gros Ventre Drainage or bison to cattle has not been documented in feed grounds) on several additional allotments in western Wyoming and the Yellowstone ecosystem the upper Gros Ventre drainage (B. Long, M. (Thorne et al. 1982; Thorne et al. 1991). Areas in Hinschberger, G. Roby, and J. Kimball. 1980. Gros the Jackson herd unit where transmission of bru­ Ventre cooperative elk study: final report 1974- cellosis from wildlife to livestock could occur are 1979. Wyoming Game and Fish Commission, un­ winter and spring ranges of elk and bison. Neither published; unpublished Bridger-Teton National species is in contact with cattle on either the Forest files; Fig. 10). Cattle herds on most private National Elk Refuge or Gros Ventre Drainage feed lands in Grand Teton National Park or adjacent to grounds or on other winter ranges in Grand Teton elk calving areas in the Bridger-Teton National National Park or the Bridger-Teton National For­ Forest likewise risk contact with B. abortus. In est. Elk occasionally feed with livestock on private allotments where grazing by livestock coincides lands in winter, primarily in the Buffalo Valley with the 25 May-15 June peak of elk parturition, east of Moran, Wyoming. This practice should be the risk of brucellosis transmission to livestock is steadfastly discouraged. greatest on federal lands (Appendix D). Where As elk and bison migrate from the National Elk grazing of allotments begins later, cattle avoid the Refuge in spring, they travel through private in­ peak parturition period of both elk and bison. holdings and cattle-grazing allotments in Grand Teton National Park (Figs. 6, 9, 10). During the Significance of Brucellosis in Jackson Hole second week of June, 2,500 head of cattle are The economic and political significance of bru­ trailed east to the Gros Ventre Mountains on a cellosis in the Jackson elk herd in a state where all stock drive trail in Grand Teton National Park livestock is certified by the U.S. Department of adjacent to the National Elk Refuge. Most elk Agriculture as free of brucellosis clearly outweighs migrate in advance of this date, but bison often the biological importance of the disease in elk. The linger in the vicinity of the drive trail near Kelly, Jackson elk herd continues to thrive and provides Wyoming. Another 800 cattle are trailed to the an annual harvest of 2,500--:3,000 animals and Blackrock-Spread Creek allotment and 235 to the unparalleled opportunities for nonconsumptive Ditch Creek allotment during early June. Cattle recreation (Smith 1991). It also provides a persist­ may encounter elk or bison in those allotments, but ent reservoir of B. abortus. the stock driveways are not significant calving Despite considerable opportunity for contact be­ areas (Fig. 10). tween cattle, elk, and bison, records (D. Woody, U.S. Cattle again trail through those stock driveways Department of Agriculture, Veterinary Services, during the last half of October. Bison often graze personal communication, 1987) that begin in 1951 the driveways in October, but the potential for show that only a single case of brucellosis in cattle transmission of brucellosis from bison or elk to that may be traceable to wild, ungulates has oc­ cattle is remote during fall. curred in Teton County, Wyoming. That case was Elk and bison calve primarily in May and June discovered during fall 1984 in a small herd of Here­ (Johnson 1951; Murie 1951; Meagher 1973; Olde­ ford cattle 11 km north of Jackson. Doubt remains meyer et al. 1993), although late-born offspring of whether this was an actual field-strain brucellosis both species have been observed during fall on the infection or a vaccination phenomenon (e.g., inad­ National Elk Refuge (Grand Teton National Park, vertent revaccination or infection with vaccine National Elk Refuge, Wyoming Game and Fish strain B. abortus). Attempts to culture organisms Department, and Bridger-Teton National Forest. from tissues of reactors were unsuccessful MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 41

(D. Woody, U.S. Department of Agriculture, Veteri­ The survival rate of male elk in 1978-84 that we nary Services, personal communication, 1987). calculated (0.63) was similar to the survival rate A 1988 infection of a cattle herd east of the (0.65) that Straley (1968) calculated of male elk in Jackson herd unit in Fremont County, Wyoming, the 1960s. Estimated survival rates during the generated a litigation for economic loss against early 1950s (x = 0.378) and late 1950s (x = 0.536) several federal and state agencies (Keiter and were not significantly different because of their Froelicher 1993). Elk and bison from Jackson Hole large standard errors (Sauer and Boyce 1983). were the alleged source of the infection. The U.S. Sauer and Boyce (1983) noted that their mean District Court of Wyoming ruled in favor of the U.S. estimates of survival may have been biased because Government in April 1992. The Wyoming Supreme they eliminated data from years when few elk were Court ruled in favor of the state in January 1993. tagged. Furthermore, our sample was from a period In both cases, convincing evidence was not pre­ when the harvest of antlerless elk was emphasized sented that either elk or bison were the source of (Appendix C). the disease in cattle. Wyoming, Montana, and Idaho are certified as free of brucellosis by the U.S. Department of Agri­ Harvests of Elk and Population culture and as such are free of mandatory testing Regulation and vaccination oflivestock. The livestock industry and the states are sensitive to any threats to that Management of the Jackson elk herd requires status. Although the probability of transmission to the cooperative efforts of several federal and state a domestic cow from a wild ungulate seems very low agencies. In 1958, the Jackson Hole Cooperative (Thorne et al. 1991; Smith and Roffe 1994), the Elk Studies Group, an interagency working group consequences are substantial for not only the live­ composed of the Wyoming Game and Fish Depart­ stock operator but for all producers who risk restric­ ment, National Park Service, U.S. Forest Service, tions of the transfer and sales of their herds. As U.S. and U.S. Fish and Wildlife Service, was formed to Department of Agriculture Veterinarian Douglas address research and management issues (Robbins Woody (personal communication, 1987) noted: et al. 1982). A primary goal of cooperative manage­ ment was to restore historic (pre-1950) summer As the goal of cattle brucellosis eradication is distributions and migrations of elk in the Jackson approached, increased attention is being herd unit (Cole 1969; National Park Service. 1986. focused on wildlife brucellosis by livestock Grand Teton National Park Natural Resources health officials. The risk of wildlife-to-cattle Management Plan and Environmental Assess­ brucellosis transmission has probably not ment, unpublished). increased, but the stakes are higher than Reasons for Hunting on the National Elk before. Refuge Hunting began on the refuge in 1943 to reduce Survival the number of wintering elk on the refuge (11,000 in winter 1942-43) and a growing summer herd (>500 in summer 1943; J. G. Griffin, 1985. Elk The mean annual survival rate of female elk in Hunting Plan National Elk Refuge. National Elk the Jackson herd unit (0.778) was not significantly Refuge, unpublished). The refuge manager sought different from the rate reported by Straley (1968) to avoid mass starvation of the wintering elk by who used the composite dynamic method reducing herd size because hay purchased annu­ (x = 0.742) and by Sauer and Boyce (1983) who used ally for feeding the elk was needed for cattle to program BROWNIE (x = 0.590 during 1958-59; help with the national shortage of meat during x = 0.679 during 1962-66). Those authors calcu­ World War II. Since the late 1960s, an increasingly lated survival rates from returned ear tags. Elk important function of the refuge hunt has been to harvests were lower in the 1950s and 1960s than assist the state of Wyoming with reducing the during the present study (Tables 14, 15). number of elk that migrate to the refuge from 42 RESOURCE PUBLICATION 199

Table 15. Number of harvested elk (Cervus elaphus) in the Jackson herd unita during 1950-1986 and the estimated number of elk in the central valley of Grand Teton National Park in subsequent summers. The estimated mean numbers of elk during 1964-1968 are significantly less (P < 0.001) than the mean numbers of elk during all later periods. The remaining mean numbers of elk do not differ from each other. Number of harvested elk in Estimated number of elk Grand Teton National Park and in the central valley of Total harvest on the National Elk Refuge Grand Teton National Park Years Management emphasis Mean ± sd Mean ± sd % of total Years Mean ± sd 1950-62 Moderate culling of elk in Grand Teton National Park 2,705 ± 917 167 ± 116 6.2 1951-63 1963-67 Stabilization of number of elk in Grand Teton National Park 3,084 ± 141 812 ± 120 23.1 1964-68 1,114 ± 25c 1968-77 Heavy harvests to control total number of elk in the Jackson herd unit 3,671 ± 1,146 667 ± 199 18.2 1969-78 1,562 ± 293c 1978-83 Reduction of the numbers of elk in Grand Teton National Park 3,356 ± 676 994 ± 106 29.6 1979-84 1,560 ± 155c 1984-86 Years of reduced harvest in the Jackson herd unit to increase numbers of elk on the National Elk Refuge but stabilize numbers in Grand Teton National Park 1,382 ± 136 507 ± 28 36.7 1985-87 1,583 ± 313d,e a5,195 Iun2 in the Snake River watershed. bNo herd size estimates (Cole 1969). c Boyce 1989. dBoyce 1989 and R. Wood, Grand Teton National Park (unpublished data). e Excluding 1986 when fog that caused poor visibility of elk resulted in a low estimate of 836 elk (R. Wood, Grand Teton National Park, personal communication).

Grand Teton National Park (J. G. Griffin, 1985. since 1950 except in 1959 and 1960. Joint state­ Elk Hunting Plan National Elk Refuge. National federal jurisdiction of elk in Grand Teton National Elk Refuge, unpublished). Hunting on the refuge Park and provisions in Public Law 81-787 are the is permitted under refuge regulations and is de­ basis for the cooperative management of hunting signed and conducted under a cooperative agree­ in the park. Cole (1969) listed the reasons and ment between the National Elk Refuge and the objectives for harvesting elk in Grand Teton Na­ Wyoming Game and Fish Department. tional Park as follows: Reasons for Hunting in Grand Teton 1. Objective: Hunt elk in the park to reduce the National Park number of early migrants to the refuge from the Grand Teton National Park herd segment. The legislation that expanded Grand Teton Na­ Reason: Many elk that summered in Grand tional Park in 1950, Public Law 81-787, provided Teton National Park migrated to the National hunting of elk in portions of the lands added to Elk Refuge in October and used forage reserved Grand Teton National Park "... when it is found for consumption in winter. necessary for the purpose of proper management 2. Objective: Harvest late-migrating elk of the Yel­ and protection of the elk." In fact, elk have been lowstone National Park herd segment to levels hunted in Grand Teton National Park every year that eventually could be controlled by hunting MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 43

outside Grand Teton National Park. National Park inversely correlated with counts of Reason: Late-migrating elk from southern Yel­ elk in the central valley in the following summers lowstone National Park that traveled through (r = -0.435, n = 22, P <0.05). Boyce (1989) did not roadless wilderness areas before crossing identify a statistically significant upward trend in Grand Teton National Park had become too the number of elk in the central valley during numerous to manage without assistance from 1963-85. However, we found that larger numbers Grand Teton National Park. of elk were counted in the central valley during 3. Objective: Restrict harvests of elk that migrate 1969-78 than during 1964-68 (t = 5.0, 13 df, east of Grand Teton National Park or summer P <0.001; Table 15). This second period of growth on national forest lands between the two parks included 4 years of record harvests (x = 4,674) in the to foster increases in those herd segments. Jackson elk herd during 1970-73, but the combined Reason: Herd segments that migrated through kills in Grand Teton National Park and in the roaded areas east of Grand Teton National Park National Elk Refuge were only 18% of all kills or summered on the more accessible national (Table 15). Based on track counts, those harvests forest lands between Yellowstone National coincided with a marked decline in the proportion Park and Grand Teton National Park had been of elk that occupied the eastern portions of the reduced to levels that no longer represented the Jackson herd unit (W. J. Barmore, 1984. Asynthesis major portion of the elk herd. of information on elk that summer in or migrate Cole (1969) stated that objectives 1 and 2 were through Grand Teton National park. Grand Teton achieved during 1963-68. Robbins et al. (1982) con­ National Park, unpublished; Boyce 1989). The long­ firmed that objective 1 was achieved. Our results term goal ofrestoring pre-1950 distributions of elk suggest that objective 2 was also achieved. Cole was abandoned to hasten reductions in the number (1969:50) further stated that progress was made of elk on feed grounds. The number of wintering elk toward achieving objective 3. He predicted "it may on the National Elk Refuge declined from 9,205 in be possible within a few years" to relegate the role 1969 and stabilized between 7,194 and 8,054 of Grand Teton National Park in the restoration of (x = 7,638) in 1971-75 (Table 4). pre-1950 distributions and migrations in the Jack­ son herd unit to a "standby status with limited Fall 1976 was extremely mild and resulted in a hunts when state programs need assistance." The late migration and the lowest harvest of elk (1,429 following evaluation indicates that Cole's (1969) elk) since 1962. The number of elk at the refuge third objective has not been achieved. grew to 8,413 during winter 1977-78. Harvest objectives from 1978 to 1983 were designed to Consequences of Harvests of Elk During control the size of the Jackson elk herd and to 1950-1985 reduce the proportion of the Grand Teton National The Grand Teton National Park herd segment Park herd segment in the herd (Smith 1985). How­ ostensibly increased during 1950-63 and 1968-78. ever, hunting seasons that extended into Decem­ Harvests from the combined Grand Teton National ber generated harvest rates of the Gros Ventre Park and National Elk Refuge hunting units dur­ Drainage, the Teton Wilderness, and Yellowstone ing 1950-62 were only 6% of the elk harvest in the National Park herd segments that were greater Jackson herd unit. As a result, the elk that sum­ than or equal to harvests of the Grand Teton mered in the Grand Teton National Park valley National Park herd segment. Numbers of elk increased from a few elk prior to 1950 (Anderson on the refuge declined to 5,010 during winter 1958) and in the early 1950s (Jepson 1952) to an 1983-84. estimated 1,430 in 1963 (Cole 1969). Mean annual harvests from the combined Increased harvests during 1963-67 in Grand Grand Teton National Park and National Elk Ref­ Teton National Park and on the National Elk Ref­ uge hunting units were highest during 1978-83 uge apparently stabilized the number of elk (Table (Tables 13, 15). Despite increased harvests and an 15). Boyce (1989) reported that after 1963, when increased proportion of females harvested in the hunting began in Hunting Units 75 and 76 in Grand park and refuge, the counts of elk in the central Teton National Park, harvests of elk in Grand Teton valley have remained stable since 1977. 44 RESOURCE PUBLICATION 199

Evaluation of Hunts in Grand Teton National Two points are cogent. Hunting in Grand Teton Park and on the National Elk Refuge National Park and the National Elk Refuge tar­ geted the Grand Teton National Park herd seg­ The Grand Teton National Park herd segment ment after 1977. Hunting-season design altered is the most difficult to harvest. About 85% of the the herd composition, but, based on estimated elk in that herd segment reside in areas presently numbers of elk in the Grand Teton National Park closed to hunting. Hunting Unit 72 represents 33% valley, park and refuge hunts did not significantly (19,838 of 59,514 ha) of the area where hunting is change the size of the Grand Teton National Park authorized by Public Law 81-787, but was opened herd segment (Tables 4 and 15). to hunting in only 1950 and 1962-67. During those 7 years, harvests in Unit 72, which is roadless, averaged only 14 elk/year (Cole 1969), which is Size and Composition of the Grand Teton insignificant in the overall management of the National Park Herd Segment Jackson elk herd. The remaining hunting units in Grand Teton National Park (75, 76, and 79) and Hunting Unit The Grand Teton National Park herd segment 77 on the National Elk Refuge are the only loca­ increased substantially after the expansion of the tions in which elk of the Grand Teton National park in 1950 and the removal oflivestock from west Park herd segment can be harvested (Fig. 3). How­ of the Snake River in 1957 (Anderson 1958; Cole ever, the distance between the Baseline Flats stag­ 1969; National Park Service, 1986. Grand Teton ing area and the southern portion of the National National Park Natural Resources Management Plan and Environmental Assessment, unpub­ Elk Refuge that is closed to hunting can be trav­ lished). Counts of the elk in the Grand Teton Na­ ersed in a few hours or overnight (as some elk do). tional Park central valley were initiated in 1963 to Mass migrations of the Grand Teton National Park monitor the number of summering elk in the most herd segment, such as the 2,700 in 1988 and 1,900 accessible and most easily investigated area of the in 1989 that arrived in the closed area of the refuge park. Based on these central valley counts, Boyce within 24 h, also thwart harvest of those elk. (1989) found an upward but statistically nonsigni­ Recognizing that some elk of the Teton Wilder­ ficant trend occurred between 1963 and 1985. Boyce ness and Yellowstone National Park herd segments (1989:49) evaluated the estimator for the central in addition to elk summering in Grand Teton Na­ valley counts and noted that "due to differential tional Park migrate through the park and refuge (lower) visibility of mature males and the increased hunting units, officials of Grand Teton National number of mature males in recent years, the total Park and the Wyoming Game and Fish Department park population may have increased more since the have consistently agreed that 40% of the total an­ mid-1960s than the trend counts indicate." We be­ nual harvest of the Jackson elk herd should be lieve another source of bias is investigators' inabil­ made in Grand Teton National Park (National Park ity to detect any redistribution of elk from the Service, 1986. Grand Teton National Park Natural relatively small area in the central valley where Resources Management Plan and Environmental counts were conducted (about 90 km2 or 8% of the Assessment, unpublished). That goal has never land surface area of Grand Teton National Park) to been achieved, not even when the harvests in the adjacent lands as a function of density-dependent park and on the refuge are combined (Table 13). effects or human and environmental influences on Since 1977, combined harvests from Grand Teton the elk or their habitat. National Park and the National Elk Refuge have Cole (1969) recognized that the central valley averaged 30.8%. Seven of28 radio-collared elk har­ count area supported the highest densities of elk in vested in Grand Teton National Park and the Na­ Grand Teton National Park. He also noted that tional Elk Refuge during this study summered out­ stochastic events, such as pine beetle control in side Grand Teton National Park. Four of the seven 1965 and 1966, seemed to alter elk distributions were killed after 30 November when 80% of the with concomitant 40-50% reductions in the central Grand Teton National Park herd segment had al­ valley count estimates of 1963. A steeper decline in ready arrived at the refuge. 1986 resulted from poor census conditions (Table 4). MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 45

These same biases and high percentages of unclass­ (2,869-4,486, 95% CI) of the elk that wintered on ified animals (.x = 64 ± 3.3%, n = 16; Table 4) could the National Elk Refuge summered in Grand Te­ likewise result in the highly improbable male-to-fe­ ton National Park. Thus, the Grand Teton Na­ male ratios during central valley counts. tional Park herd segment composed one-third of The counts of elk in the central valley are not the targeted number of elk in the Jackson herd censuses of elk on lands east of State Highway unit (3,700 of 11,000 = 33.6%). 26/89/187 in Grand Teton National Park, on either Throughout this study, removals of elk of the side of Jackson Lake, along the Snake River ripar­ Grand Teton National Park herd segment during ian corridor, or north, south, and west of the inside fall hunting seasons averaged 17%. Therefore, an Park Road (Moose to Jenny Lake to Jackson Lake; estimated 4,460 (± 981, 95% CI) elk summered Fig. 1). Any changes in elk numbers in those areas in Grand Teton National Park prior to the fall would not be detected. Replicate counts by helicop­ harvests. ter were initiated in 1990. They census elk in a larger area and should clarify the precision and accuracy of the valley count estimator (S. Cain and Population Regulation B. Smith, in preparation). Cole's (1969) estimate of 2,800 elk in Grand Recruitment and Survival Teton National Park during summer 1964 from a Based on observed density-dependent recruit­ combination of ground, aerial, and track counts ment, Boyce (1989) predicted that the number of was the first attempt of a census in an area larger elk in the Grand Teton National Park central than the central valley of Grand Teton National valley (in the limited area where the valley count Park. Houston (1968) estimated the size of the estimator is applied) would stabilize at about Grand Teton National Park herd segment as 3,250 2,000 animals in the absence of hunting. He dis­ during fall 1968. cussed an experimental suspension of hunting in The best direct count of the Grand Teton Na­ Grand Teton National Park as proposed in the tional Park herd segment was made in 1986. Dur­ Grand Teton National Park Natural Resources ing an aerial survey on 12 November 1986, Wyo­ Management Plan (National Park Service, 1986. ming Game and Fish Department biologists Grand Teton National Park Natural Resources counted 685 elk along the west side of Jackson Management Plan and Environmental Assess­ Lake and another 724 in the Grand Teton National ment, unpublished) to determine whether elk in Park valley. Back-tracking and track count data the Grand Teton National Park valley would self­ indicated that these were elk of the Grand Teton regulate their numbers. National Park herd segment. On the same day, Density-dependent regulation of ungulate popu­ 2,296 elk were counted and classified on the Na­ lations is generally a function of the combined tional Elk Refuge. These were primarily elk of the effects of environmental conditions and reduced Grand Teton National Park herd segment because forage quality and quantity on natality, survival, the earliest migrants to the National Elk Refuge and dispersal (Sinclair 1977; Clutton-Brock et al. are from Grand Teton National Park (Cole 1969; 1982). These effects are usually manifested in elk Smith 1985; this study). The total of 3,705 elk does of the Rocky Mountain West during winter and not include remaining elk in the eastern portions spring months (Flook 1970; Craighead et al. 1973; of Grand Teton National Park or animals that had Houston 1982). been harvested during the Grand Teton National Winter feeding has ameliorated the nutritional Park and National Elk Refuge hunting seasons stress and limited over-winter mortality of elk to that opened on 25 October. an average of 1.4% annually on the National Elk A corroborative estimate of the size of the Refuge during the past 20 years (Boyce 1989; Grand Teton National Park herd segment was Smith and Wilbrecht 1990). Since 1960, the per­ derived from the summer distributions of the ra­ centage of winter mortality has not correlated dio-collared elk. From 1978 to 1982, an average of with the number of elk that wintered on the refuge 7,630 elk wintered on the National Elk Refuge. An (r = -0.226, n = 28, P >0.05; Smith 1985). average of 48.2% of the radio-collared elk or 3,678 Boyce (1989) found evidence of density-dependent 46 RESOURCE PUBLICATION 199

recruitment among the elk that wintered on the National Elk Refuge during some summers. He National Elk Refuge in only 5 years since 1940 postulated that in some years a substantial frac­ when he estimated feed-ground attendance by elk tion of yearlings do not migrate. The large number should have been highest because winter severity of yearlings that have summered in Grand Teton was greatest. National Park and on the refuge since 1969 sup­ Free-ranging populations of large mammals port Martinka's contention. The natal summer generally respond in a density-dependent fashion ranges of these nonmigratory yearlings is un­ to harvests that hold the populations below their known. Failure to return to natal summer ranges ecological carrying capacity (Eberhardt 1977; outside Grand Teton National Park would add to Caughley 1979; McCullough 1984). Boyce (1989) the annual rate of increase of the Grand Teton attributed a significant increase in young-to-fe­ National Park herd segment. male-elk ratios after 1976 in Grand Teton Na­ tional Park to an increased harvest of antlerless elk. Whether the source of the density-dependent Restoration of Historic Distributions of recruitment was fecundity or survival of young Elk were uncertain. Similar data for the other herd segments are not available. The restoration of pre-1950 distributions and Because of the nonlinearity in productivity migrations of elk east of Grand Teton National Park curves of large mammals, the greatest density-de­ remains an important objective of state and federal pendent change among free-ranging populations is agencies because of the recreational, aesthetic, eco­ expected near carrying capacity (Fowler 1981). The nomic, and ecological benefits of conserving elk relation between densities of elk in Grand Teton throughout all summer ranges in the national for­ National Park and the ecological carrying capacity est and the national parks. A significant decline in (K; Caughley 1979) of these animals' summer range the number of elk migrating through the eastern is unclear. Boyce (1989) asserted that harvests portion of the Jackson herd unit since 1950 has been since the mid-1960s have kept the number of elk in blamed on high harvest rates there (Boyce 1989). the Grand Teton National Park central valley below The lower harvest rates of radio-collared elk along K and possibly nearer a density that provides a eastern migration corridors offer encouragement maximum sustained yield. Given the ameliorating that harvest strategies since 1978 may restore the role of winter feeding on density effects in all herd elk of the eastern Yellowstone National Park and segments, investigations of differences in fecundity, the Teton Wilderness herd segments to pre-1950 survival of young, andjuvenile dispersal among the densities. Heavy antlerless harvests from 1977 to herd segments may clarify the role of density-de­ 1983 skewed sex ratios and probably dampened the pendence in the Jackson elk herd. growth rate of the Grand Teton National Park herd Dispersal segment. But the overall size of that herd segment seemingly was not reduced. The emphasis on an­ Dispersal can be an important regulating tlerless harvests was abandoned from 1984 to 1986. mechanism of populations that is most pro­ We predict that (1) in the absence of effective strate­ nounced among males of polygynous mammals gies to harvest elk that summer in and migrate (Dobson 1982). At present densities, we found that through Grand Teton National Park, restoration of adult radio-collared elk are not dispersing from the pre-1950 distribution of elk in the Jackson herd Grand Teton National Park. Houston (1982) and unit will not occur and (2) the Grand Teton National Clutton-Brock et al. (1982) reported density-de­ Park herd segment will continue to compose at least pendent recruitment and mortality but found lit­ half of the elk that winter on the National Elk tle evidence ofregulation by dispersal ofhigh-den­ Refuge. sity populations of elk and red deer that were not We submit that experimental suspension of hunted. hunting in Grand Teton National Park, as pro­ Martinka (1969) found higher-than-expected posed in the Grand Teton Natural Resource Man­ numbers of yearling males and females in the agement Plan (National Park Service, 1986. Grand Teton National Park valley and on the Grand Teton National Park Natural Resources MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 4 7

Management Plan and Environmental Assess­ To achieve Cole's (1969) third objective through ment, unpublished), is a high-risk proposition. herd removals will require changes in two areas Once suspended, public sentiment may prohibit of management: (1) establishing specific objec­ the reinstatement of hunting in Grand Teton Na­ tives for the spatial distribution of the number of tional Park (Boyce 1989). Without a hunt in Grand elk on summer ranges; and (2) adopting new Teton National Park, a burgeoning elk herd may strategies to achieve elk removals that favor elk require massive reductions by agency personnel that summer in areas beyond Grand Teton Na­ on the National Elk Refuge. In 1935, refuge and tional Park and that migrate east of Grand Teton state of Wyoming personnel culled 500 elk on the National Park. National Elk Refuge to help control herd size. This The other alternative is to terminate hunting one-time cull was deemed unacceptable by the on the National Elk Refuge and in Grand Teton public. Extensive culling of elk by the National National Park and test the hypothesis that den­ Park Service during the 1950s and 1960s was sity-dependent regulation stabilizes the Grand Te­ abandoned in 1969 as inappropriate management ton National Park herd segment at or below its in Yellowstone National Park (Houston 1982). current size. A yield curve generated for elk that Annual harvests on the National Elk Refuge summer in the Grand Teton National Park central have averaged about 30% of the combined Grand valley contains only one data point near the pre­ Teton National Park and refuge harvests dicted K (Boyce 1989), which brings into question (J. G. Griffin, 1985. Elk Hunting Plan National its reliability. More importantly, that model per­ Elk Refuge. National Elk Refuge, unpublished). tains only to a small part of the summer range of Without hunting on the National Elk Refuge and the Grand Teton National Park herd segment. in Grand Teton National Park to remove elk of the Predicting K is far from a precise science Grand Teton National Park herd segment, the (Caughley 1976). In the absence of supplemental other herd segments require harvests that are feeding, elk and other ungulate populations (Tan­ higher than their annual rates of increase to ner 1966; Caughley 1970; Fowler 1981) peri­ hold numbers of elk on the refuge below 7,500. odically exceed Kby a considerable margin as they Alternatively, accepting larger numbers of elk oscillate in dynamic equilibrium with their envi­ on feed grounds would escalate economic and eco­ ronment. Rather than debate whether we can pre­ logical costs that are contrary to sound resource dict the carrying capacity of Grand Teton National stewardship. Park and whether that number is acceptable for One can argue that the humans' role in corrupt­ managing the Jackson elk herd in accordance with ing an evolved ecosystem has necessitated humans' current policy constraints, we must first decide if role in manipulating the number of elk. The usur­ human non-interference in ecological processes is pation of historic winter ranges, removal of large appropriate and practical in Jackson Hole. By predators, suppression of wildfire, establishment of definition, natural regulation would require dras­ the Yellowstone National Park and Grand Teton tic reduction, if not elimination, of the supplemen­ National Park ecological reserves in portions of the tal winter feeding of elk. Cayot et al. (1979) cor­ ecosystem, and selective modification of habitats rectly stated that natural regulation of elk outside those reserves through timber harvesting, populations is valid management in national road construction, and residential and agricultural parks with ecologically complete habitats. Pre­ encroachment have played a part in the legacy of sumptions about how or what may naturally regu­ Jackson Hole and its resilient elk herd. To mitigate late the elk population under prevailing condi­ these failings and maintain more than a remnant tions in Jackson Hole stretch the limits of of the former herd, supplemental feeding was en­ ecological theory. dorsed and institutionalized by the federal and The elimination of hunting in Grand Teton Na­ state governments (Robbins et al. 1982). As an tional Park alone would not restore a naturally ironic consequence of its success, herd reductions functioning ecosystem. Any lessons to be learned through public hunts became necessary in from the exercise would be confounded by other Grand Teton National Park and on the National Elk human interference and have questionable appli­ Refuge. cation elsewhere. Results from testing hypotheses 48 RESOURCE PUBLICATION 199

about natural regulation are probably inconclu­ radio-tagging neonates in Grand Teton Na­ sive unless research is conducted in truly intact or tional Park and in the Bridger-Teton National restored ecosystems (Peek 1980). Forest to investigate regulation of herd-seg­ ment sizes on summer ranges with high and low densities of elk. This research would ad­ Recommended Research dress the premise for terminating the hunting of elk in Grand Teton National Park. 4. Results from the recommended studies should 1. The accuracy and precision of classification be incorporated into population-growth models counts of elk in the Grand Teton National Park for the Grand Teton National Park herd seg­ valley should be verified by replication. Ongoing ment and, if possible, for the other herd seg­ research (S. Cain and B. Smith, in preparation) ments. These models may help predict the out­ indicates that the high male-to-female-elk ratios comes of alternative management of the Jack­ obtained from the central valley classification son elk herd. counts are inflated. Two factors bias those clas­ sifications: (1) the geographic area of the counts is perennially favored by males, and (2) the high Recommended Management percentage (64 ± 3.3%, n = 16) of unclassified animals that are largely female-young groups among all observed elk. Preliminary results in­ Population Management dicate that counts from helicopters are superior to the traditional counts from the ground for We offer four distinct options for the manage­ estimating the herd size and composition. ment of the Jackson elk herd on the assumptions 2. After objectives for herd segments are estab­ that (1) the current objective ofll,000 wintering elk lished (see Management Recommendations), in the Jackson herd unit remains in effect, and (2) systematic aerial trend censuses should be im­ the responsible state and federal agencies continue plemented to monitor progress toward those to manage the herd for a variety of consumptive and objectives. Annual track counts enable wildlife nonconsumptive values. managers to assess the relative numbers of elk Three options embrace the goal of restoring the that migrate from eastern and western por­ pre-1950 distributions of elk in summer. That goal tions of the Jackson herd unit. However, elk requires reducing the Grand Teton National Park from two or three herd segments cross most herd segment by about half and concomitantly track-count routes. increasing the number of elk on national forest Previous aerial counts of elk on summer ranges lands and in eastern Yellowstone National Park. did not account for visibility biases and were Objectives for the desired size and age-sex compo­ discontinued because of the variability of the sition of each herd segment would be established results (Anderson 1958). Recently developed and would drive management decisions. The models for aerial winter counts of elk can cor­ fourth option would not establish an objective for rect inherent visibility biases (Samuel et al. the size of the Grand Teton National Park herd 1987). Advancements in aerial census tech­ segment. niques and data analysis (Caughley 1974; Option 1. Refine current hunting regulations to Caughley et al.1976; Samuel 1984) should be achieve management objectives. Annual harvests adopted to measure the sizes and compositions of antlerless elk are necessary to restrain growth of herd segments in summer. The extensive of the Grand Teton National Park herd segment. forest fires of 1988 in Yellowstone National The opportunity to harvest elk of the Grand Teton Park and in the Teton Wilderness should mark­ National Park herd segment is influenced tempo­ edly improve the visibility of elk in formerly rally by environmental conditions that initiate heavily forested areas. migration to the National Elk Refuge and by the 3. Recruitment, survival, and dispersal ofjuvenile increasing abundance of migrants from Yellow­ elk should be determined by capturing and stone National Park and the Teton Wilderness in MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 49

Hunting Units 75, 76, 77, and 79 as the migration winter distributions and forage use by elk (reduc­ progresses. Offer the purchase of a second permit ing the need for artificial feeding) by closing or for antlerless elk to hunters in Hunting Unit 77 shortening the hunting seasons on potential win­ and possibly in Hunting Units 75 and 76 to in­ ter range in Hunting Units 75-77. Although it is crease harvests while elk of the Grand Teton Na­ an ecologically rational approach to the elk man­ tional Park herd segment are migrating to the agement issue, this alternative will be unaccept­ National Elk Refuge. Hunting seasons should able to members of the American public that object close before 23 November in Hunting Unit 77 and to hunting in national parks. before 1 December in Hunting Units 75, 76, and Option 3. Conduct periodic direct reductions of 79-earlier if targeted herd segments have passed early migrants to the National Elk Refuge to reduce through the areas and later only if elk migrations the Grand Teton National Park herd segment. Be­ are greatly delayed by mild weather. Open the cause the Jackson elk herd is infected with brucel­ hunting seasons in Hunting Units 71, 79, and 81 losis, live elk cannot leave Teton County unless on 1 October. Concurrent opening dates may dis­ consigned for slaughter or first quarantined and perse some elk from Grand Teton National Park tested for disease (Wyoming State statute 11-19- east into adjacent national forest lands where they 101; Legislative Service Office 1989). Suitable habi­ may be harvested. tat elsewhere in the county is occupied, and the elk To reduce harvests of elk in the eastern portion herds exceed desired numbers. of the Jackson herd unit, move the opening of hunt­ Direct reductions will elicit disapproval from the ing season in Hunting Unit 70 from 10 September public. An alternative is the reduction or periodic to 1 October. Close the hunting season in Hunting elimination of winter-feed rations. Elk from all herd Unit 81 on 10 November to protect elk migrating to segments would suffer nutritional stress and in­ winter ranges in the Gros Ventre drainage and on creased mortality. This is contrary to current man­ the National Elk Refuge. Practices that enhance agement policies of the National Elk Refuge and the habitat effectiveness (Thomas 1979) for elk along Wyoming Game and Fish Department. Remotely migration corridors (including the maintenance of delivered sterilants or abortifacients may offer a security cover and seasonal road closures) and on more humane alternative to direct reductions or identified calving areas and winter ranges should deprivation of food. receive high priority from land managers. Option 4. Experimentally terminate hunting in Option 2. Amend Public Law 787-81 to permit Grand Teton National Park and allow density-de­ hunting in Grand Teton National Park west of the pendent factors to regulate the Grand Teton Na­ Snake River where most of the elk in Grand Teton tional Park herd segment. This is the preferred National Park summer. Growth of the Grand Te­ alternative in the Grand Teton National Park Re­ ton National Park herd segment has resulted from source Management Plan for management of elk low winter mortality and inadequate harvests in (National Park Service, 1986. Grand Teton Na­ the limited geographical areas in which elk can be tional Park Natural Resources Management Plan hunted. Periodic harvests of elk west of the Snake and Environmental Assessment, unpublished). River would achieve (1) sufficient harvests of elk Although this alternative is appealing, insuffi­ of the Grand Teton National Park herd segment cient evidence suggests that the herd segment will before they mingle with elk from other herd seg­ decline or even stabilize at its present size with ments; (2) desired sex ratios of elk in Grand Teton continuance of winter feeding on the National Elk National Park; (3) reduced harvest pressure, ex­ Refuge. cept when and if needed, on the elk of the Gros In addition to these four alternatives, we see Ventre drainage, Teton Wilderness, and Yellow­ merit in management that requires more extreme stone National Park herd segments that migrate departures from the current management philoso­ to the National Elk Refuge by diminishing or phy. Winter feeding of elk should not be accepted as eliminating the need for hunting in Hunting Units the necessary norm for the Jackson elk herd. It is 75, 76, and 77; (4) harvests of elk in northwest costly, increases disease prevalence, and causes Grand Teton National Park without reopening range damage (Thorne et al. 1979; Boyce 1989; Hunting Unit 72 to hunting; and (5) improved Smith and Roffe 1994). Improving the distribution 50 RESOURCE PUBLICATION 199

of elk in winter throughout existing and potential strongly discourage intentional and incidental feed­ winter ranges and substantially reducing winter ing of elk or bison together with livestock. Thirdly, feeding are worthy management goals. Ongoing decreasing the dependence of the Jackson elk herd range improvements (particularly prescribed burn­ on supplemental feeding will reduce winter concen­ ing, seeding and irrigation of refuge and forest trations and disease transmission. rangelands, and reduction of livestock grazing on AB an integral component of the greater Yellow­ critical winter range in the Gros Ventre drainage) stone area, the Jackson elk herd tests our commit­ are collectively enhancing the carrying capacities ment to the ideals of maintaining wild places and wild for free-ranging elk. These practices, conservation things. AB a priceless slice of our wildlife heritage, the easements, and acquisitions of available privately elk herd is a microcosm of the challenges elk-and owned habitat are vital for a secure future of the people-will face throughout the coming years. Jackson elk herd. AB these changes gradually shift winter distributions from the National Elk Refuge to winter ranges in the Bridger-Teton National For­ est and in Grand Teton National Park, the propor­ Acknowledgments tion of elk of the Jackson elk herd that summer in Grand Teton National Park should decline. This research was funded by the U.S. Fish and This redistribution of elk in winter could be Wildlife Service; additional funding for flight time achieved without lowering the size of the Jackson was contributed by the Safari Club International. elk herd below the current goal of 11,000 elk. About G. E. Roby of the Wyoming Game and Fish Depart­ 14,000-15,000 elk are currently wintering in the ment and D. E. Griffel of the Bridger-Teton Na­ Jackson herd unit and over 4,000 of those are not tional Forest helped immobilize elk. G. E. Roby, on feed grounds (D. Moody and T. Toman, Wyoming T. L. Toman, E.T. Thorne, and D. A. Crawford of Game and Fish Department, personal communica­ the Wyoming Game and Fish Department and tion, 1993). To formalize a change in the number of staff of the National Elk Refuge assisted with elk that are fed in winter, the state-federal agree­ trapping. F. A. Gordon, Bridger-Teton National ment that endorses the maximum number of elk that winter on the National Elk Refuge should be Forest, R. P. Wood and S. L. Cain, Grand Teton changed from 7,500 to 5,500-as suggested by National Park, provided data from agency files. Murie (1951)-or fewer animals. One or more of the G. C. White assisted with statistical analyses of methods listed in management alternatives 1-3 are survival rates. Numerous seasonal employees and required to achieve an initial reduction to 5,500 or volunteers at the National Elk Refuge assisted fewer wintering elk on the refuge. Required annual with collecting and cataloguing field data. A spe­ harvests and supplemental feeding will decline in cial thanks is due M. J. Heisler for radio monitor­ future years as the number of elk that winter on the ing of elk and many other technical contributions refuge declines. to the research. D. E. Biggins, M. S. Boyce, D. B. Houston, and L. L. McDonald, provided helpful reviews of an Brucellosis Management earlier draft of the manuscript.

Long et al. (B. Long, M. Hinschberger, G. Roby, and J. Kimball, 1980. Gros Ventre cooperative elk Cited Literature study: final report 1974-1979. Wyoming Game and Fish Commission, Cheyenne. 192 pp.) advised de­ Adams, A. W. 1982. Migration. Pages 301-321 in J. W. laying the initiation of grazing on calving areas Thomas and D. E. Toweill, editors. Elk ofNorth Amer­ until late June or July to avoid contact by cattle ica: ecology and management. Stackpole Books, Har­ with elk fetuses and afterbirths. We agree and also risburg, Penn. recommend continued, resolute vaccination of Altmann, M. 1952. Social behavior of elk, Cervus ca­ young of all cattle in Teton and other counties with nadensis nelsoni, in the Jackson Hole area of Wyo­ infected herds of elk and bison. Secondly, we ming. Behavior 4:116-143. MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 51

Anderson, C. C. 1958. The elk of Jackson Hole: a review Craighead, J. J., G. Atwell, and B. W. O'Gara. 1972. Elk of Jackson elk studies. Wyoming Game and Fish migrations in and near Yellowstone National Park. Commission, Bulletin 10, Cheyenne, Wyo. 184 pp. Wildlife Monograph 29. 48 pp. Bendt, R. H. 1962. The Jackson Hole elk herds in Craighead, J. J., F. C. Craighead, Jr., R. L. Ruff, and Yellowstone and Grand Teton National parks. Trans­ B. W. O'Gara. 1973. Home ranges and activity pat­ actions of the North American Wildlife Conference terns of non-migratory elk of the Madison drainage 27:191-201. as determined by biotelemetry. Wildlife Monograph Blunt, F. M. 1950. Migration study of the Jackson Hole 33. 50 pp. elk herd. Wyoming Wildlife 14(2):25-32. Davis D.S., J. W. Templeton, T. A. Ficht, J. D. Williams, J. D. Kopec, and L.G. Adams. 1988. Brucella abortus Boyce, M. S. 1989. Elk management in North America: in captive bison. 1. Serology, bacteriology, pathogene­ the Jackson herd. Cambridge Press, Cambridge, sis, and interspecific transmission to susceptible Bos United Kingdom. 306 pp. taurus heifers. Abstract only. Wildlife Disease Asso­ Boyce, M. S. 1991. Migratory behavior and management ciation Conference, Athens, Ga. August 1988. of elk (Cervus elaphus). Applied Animal Behavior Davis D.S., J. W. Templeton, T. A. Ficht, J. D. Huber, R. D. Science 29:239-250. Angus, and L. G. Adams. 1991. Brucella abortus in Brazada, A. R. 1953. Elk migration patterns and some bison. II. Evaluation of Strain 19 vaccination of preg­ of the factors affecting movements in the Gallatin nant cows. Journal of Wildlife Diseases 27:258-264. River drainage. Journal of Wildlife Management Dexter, D. 1984. Sixty minutes and the Teton elk. Wyo­ 17:9-29. ming Wildlife 48:3. Brown, R. C. 1947. The Jackson Hole elk herd. Wyoming Dobson, F. S. 1982. Competition for mates and predomi­ Wildlife 11(12):4-11, 29-32. nant juvenile male dispersal in mammals. Animal Byers, C.R., R. K. Steinhorst, and P.R. Krausman. 1984. Behavior 30:1183-1192. Clarification of a technique for analysis ofutilization­ Eberhardt, L. L. 1977. "Optimal" management policies availability data. Journal of Wildlife Management for marine mammals. Wildlife Society Bulletin 5:162-169. 49:1050-1053. Flook, D. R. 1970. A study of sex differential in the Caughley, G. 1970. Eruption of ungulate populations survival of wapiti. Canadian Wildlife Service Report with emphasis on Himalayan thar in New Zealand. 11, Ottawa, Ontario. 71 pp. Ecology 51:53-72. Fowler, C. W. 1981. Comparative population dynamics Caughley, G. 1974. Bias in aerial survey. Journal of in large mammals. Pages 437-455 in C. W. Fowler Wildlife Management 38:921-933. and T. D. Smith, editors. Dynamics oflarge mammal Caughley, G. 1976. Wildlife management and the dy­ populations. John Wiley and Sons, New York. namics of ungulate populations. Pages 183-246 in Fowler, C. W. 1981. Density dependence as related to life T. H. Coaker, editor. Applied biology, Volume 1, Aca­ history strategy. Ecology 62:602-610. demic Press, New York. Greer, K. R., and W.W. Hawkins, Jr. 1967. Determining Caughley, G. 1979. What is this thing called carrying pregnancy in elk by rectal palpation. Journal of Wild­ capacity? Pages 2-8 in M. S. Boyce, and L. D. Hay­ life Management 31:145-149. den-Wing, editors. North American elk: ecology, be­ Hershey, T. J., and T. A. Leege. 1982. Elk movement and havior and management. University of Wyoming, habitat use on a managed forest in north-central Laramie. Idaho. Idaho Department of Fish and Game, Wildlife Caughley, G., R. Sinclair, and D. Scott-Kemmis. 1976. Bulletin 10, Boise. 24 pp. Experiments in aerial survey. Journal of Wildlife Houston, D. B. 1968. The Shiras moose in Jackson Hole, Wyoming. Technical Bulletin Number 1. Grand Teton Management 40:290-300. Natural History Association, Moose Wyo. 110 pp. Cayot, L. J., J. Prukop, and D.R. Smith. 1979. Zootic Houston, D. B. 1982. The northern Yellowstone Elk. climax vegetation and natural regulation of elk in Macmillan Publishing Company, New York. 4 74 pp. Yellowstone National Park. Wildlife Society Bulletin Jepson, C. E. 1952. Report on the elk survey and migra­ 7:163-169. tion studies, 1952. National Park Service, Grand Te­ Clutton-Brock, T. H., F. E. Guinness, and S. D. Albon. ton National Park, Moose, Wyo. 1982. Red deer: behavior and ecology of two sexes. Johnson, D. E. 1951. Biology of the elk calf, Cervus University of Chicago Press, Ill. 378 pp. canadensis nelsoni. Journal of Wildlife Management Cole, G. F. 1969. The elk of Grand Teton and southern 15:396-410. Yellowstone National parks. National Park Service Johnson, R. 1976. Elk and logging activities. Wyoming Research Report GRTE-N-1, Washington, D.C. 80 pp. Wildlife 40:8-11. 52 RESOURCE PUBLICATION 199

Knight, R. R. 1970. The Sun River elk herd. Wildlife ica: ecology and management. Stackpole Books, Har­ Monograph 23. 66 pp. risburg, Penn. Keiter, R. B., and P.H. Froelicher. 1993. Bison, brucello­ Neu, C. W., C.R. Byers, andJ. M. Peek.1974.Atechnique sis, and law in the Greater Yellowstone Ecosystem. for analysis of utilization-availability data. Journal of Land and Water Law Review. Vol. 28. No. 1. Univer­ Wildlife Management 38:541-545. sity College of Law, Laramie. 75 pp. Oldemeyer, J. L., R. L. Robbins, and B. L. Smith. 1993. Kuck, L., G. L. Hompland, and E. H. Merrill. 1985. Elk Effect offeeding level on elk weights and reproductive calf response to simulated mine disturbance in success at the National Elk Refuge. In R. Callas, D. southeast Idaho. Journal of Wildlife Management Koch, and E. Loft, editors. Western States and Prov­ 49:751-757. inces Elk Workshop, California Fish and Game De­ Legislative Service Office. 1989. Wyoming statutes an­ partment, Eureka, Calif. notated 1977 republished edition. The Michie Com­ Peek, J.M. 1980. Natural regulation ofungulates (What pany, Charlottesville, N.C. constitutes a real wilderness?). Wildlife Society Bul­ Love, J. D., and J.C. Reed, Jr. 1968. Creation of the Teton letin 8:217-227. Landscape. Grand Teton Natural History Associa­ Preble, E. A. 1911. Report on condition of elk in Jackson tion, Moose, Wyo. Hole, Wyoming in 1911. U.S. Department of Agricul­ Manthei, C. A. 1952. Evaluation ofvaccinal methods and ture, Bureau of Biological Survey Bulletin 40, Wash­ doses of Brucella abortus Strain 19. Proceedings of ington, D.C. 25 pp. the U.S. Livestock Sanitary Association 56:115-125. Quimby, D. C., and J.E. Gaab. 1957. Mandibular denti­ tion as an age indicator in Rocky Mountain elk. Jour­ Manthei, C. A. 1959. Summary of controlled research with Strain 19. Proceedings of the U.S. Livestock nal of Wildlife Management 21:435-451. Sanitary Association 63:91-97. Rippe, D. D., and R. L. Rayburn. 1981. Land use and big game population trends in Wyoming. U.S. Fish and Martinka, C. J. 1969. Population ecology of summer Wildlife Service, Biological Services Program W/0 resident elk in Jackson Hole, Wyoming. Journal of CRAM-81/W22. Fort Collins, Colo. 27 pp. Wildlife Management 33:465-481. Robbins, R. L., D. E. Redfearn, and C. P. Stone. 1982. McCullough, D. R. 1984. Lessons from the George Re­ Refuges and elk management. Pages 4 79-507 in J. W. serve, Michigan. Pages 211-242 in L. K. Halls, editor. Thomas and D. E. Toweill, editors. Elk ofNorth Amer­ White-tailed deer: ecology and management. Stack­ ica: ecology and management. Stackpole Books, Har­ pole Books, Harrisburg, Penn. risburg, Penn. Meagher, M. M. 1973. The bison ofYellowstone National Roelle, J. E., and G. T. Auble. 1983. Resource develop­ Park. National Park Service Scientific Monograph 1. ment and management in Jackson Hole, Wyoming. 161 pp. Pages 303-311 in W. K. Laurenroth, G. V. Skogerboe, Mitchell, B. 1963. Growth layers in dental cement for and M. Flug, editors. Development in environmental determining age of red deer. Journal of Animal Ecol­ modelling 5: Analysis of ecological systems: state of ogy 36:279-293. the art in ecological modelling. Elsevier, N.Y. Moen, A. N. 1973. Wildlife ecology, W. H. Freeman and Rudd, W. J., A. L. Ward, and L. L. Irwin. 1983. Do split Co., San Francisco. 458 pp. hunting seasons influence elk migrations from Yel­ Morgantini, L. E., and R. J. Hudson. 1988. Migratory lowstone National Park? Wildlife Society Bulletin patterns of the wapiti, Ceruus elaphus, in Banff Na­ 11:328-331. tional Park, Alberta. Canadian Field Naturalist Samuel, M. D. 1984. An evaluation of elk sightability in 102:12-19. northcentral Idaho with application to aerial census Murie, 0. J. 1951. The elk of North America. Stackpole and herd classification counts. Ph.D. thesis, Univer­ Books, Harrisburg, Penn. 376 pp. sity of Idaho, Moscow. 118 pp. Murie, M. E., and 0. J. Murie. 1966. Wapiti wilderness. Samuel, M. D., E. 0. Garton, M. W. Schlegel, and R. G. Alfred A. Knopf Publishing, New York. 302 pp. Carson. 1987. Visibility bias during aerial surveys of National Oceanic and Atmospheric Administration. elk in northcentral Idaho. Journal of Wildlife Man­ 1982. Monthly station normals of temperature, pre­ agement 51:622-630. cipitation, and heating and cooling degree days 1961- Sauer, J. R. 1980. The population ecology of wapiti in 1990. Climatography of the United States No. 81. northwestern Wyoming. M.S. thesis, University of National Climatic Data Center, Asheville, N.C. 27 pp. Wyoming, Laramie. 43 pp. Nelson, J. R., and T. A. Leege. 1982. Nutritional require­ Sauer, J. R, and M. S. Boyce. 1983. Density dependence ments and food habits. Pages 323-368 in J. W. and survival of elk in northwest Wyoming. Journal of Thomas and D. E. Toweill, editors. Elk of North Amer- Wildlife Management 47:31-37. MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 53

Seton, E. T. 1927. Lives of game animals. Vol. 3, part 1. Tanner, J. T. 1966. Effects ofpopulation density on growth Doubleday, Page, and Co, Garden City, N.Y. 412 pp. rates of animal populations. Ecology 4 7 :733-745. Sinclair, A. R. E. 1977. The African buffalo-a study of Taylor, D. D., E. B. Bradley, and M. M. Martin. 1982. The resource limitation of populations. University of Chi­ outfitting industry in Teton County: its clientele and cago Press, Ill. 355 pp. economic importance. Wyoming Agricultural Exten­ Skovlin, J. M. 1982. Habitat requirements and evalu­ sion Service Bulletin B-793. University of Wyoming, ations. Pages 365-413 in J. W. Thomas and D. E. Laramie. 19 pp. Toweill, editors. Elk of North America: ecology and Thomas, E., D. Crowe, and L. Kruckenberg. 1984. Man­ management. Stackpole Books, Harrisburg, Penn. aging Teton elk. Wyoming Wildlife 48(5):18-21. Skovlin, J.M., P. J. Edgerton, and R. W. Harris. 1968. Thomas, J. W., editor. 1979. Wildlife habitats in managed The influence of cattle management on deer and elk. forests: the Blue Mountains of Oregon and Washing­ Transactions of the North American Wildlife and ton. U.S.D.A. Forest Service, Agriculture Handbook Natural Resources Conference 33:169-181. 553. U.S. Government Printing Office, Washington, D.C. 512 pp. Smith, B. L. 1985. Scabies and elk mortalities on the Thorne, E. T., R. E. Dean, and W. G. Hepworth. 1976. National Elk Refuge. Pages 180-194 in R. W. Nelson, Nutrition during gestation in relation to successful editor. Proceedings of the Western States and Prov­ reproduction in elk. Journal of Wildlife Management inces Elk Workshop, Alberta Fish and Wildlife Divi­ 40:330-335. sion, Edmonton. Thorne, E. T., J. K. Morton, and W. C. Ray. 1979. Brucel­ Smith, B. L. 1991. Jackson: the big herds. Bugle 8:48-58. losis, its effect and impact on elk in western Wyoming. Smith, B. L. 1994. Out-of-season births of elk calves in Pages 212-220 in M. S. Boyce and L. D. Hayden­ Wyoming. The Prairie Naturalist. In press. Wing, editors. North American elk: ecology, behavior Smith, B. L., and J. W. Wilbrecht. 1990. Supplemental and management. University of Wyoming, Laramie. winter feeding of elk on the National Elk Refuge, Thorne, E. T., N. Kingston, W. R. Jolley, and R. Wyoming. Pages 30-41 in M. Zahn, J. Pierce, and R. Bergstrom. 1982. Diseases of wildlife in Wyoming. Johnson, editors. Proceedings of the Western States Wyoming Game and Fish Department, Cheyenne. and Provinces Elk Workshop, Washington Depart­ 353 pp. ment of Wildlife, Olympia, Wash. Thorne, E.T., M. Meagher, and R. Hillman. 1991. Bru­ Smith, B. L., and T. J. Roffe. 1994. Diseases among elk cellosis in free-ranging bison: three perspectives. of the Yellowstone ecosystem, U.S.A. Third Interna­ Pages 275-287 in R. B. Keiter and M. S. Boyce, tional Wildlife Ranching Symposium, Pretoria, South editors. The Greater Yellowstone ecosystem: redefin­ Africa. In press. ing American wilderness heritage. Yale University Sokal, R. R., and F. J. Rohlf. 1981. Biometry. 2nd ed. Press, New Haven, Conn. W. H. Freeman and Company, New York. 859 pp. Toman, T. 1987. Tangled trails in Jackson Hole. Wyo­ Straley, J. H. 1968. Population analysis of ear-tagged ming Wildlife 51:14-19. elk. Proceedings of the Western Association of State White, G. C. 1983. Numerical estimation of survival Game and Fish Commissioners 48:152-160. rates from band-recovery and biotelemetry data. Sweeney, J. M., and H. W. Steinhoff. 1976. Elk move­ Journal of Wildlife Management 47:716-728. ments and calving as related to snow cover. Pages Williams, E. S., E. T. Thorne, S. L. Anderson, and J. D. 415-436 in H. W. Steinhoff and J. D. Ives, editors. Herriges. 1993. Brucellosis in free-ranging bison (Bi­ Ecological impacts of snowpack augmentation in the son bison) from Teton County, Wyoming. Journal of San Juan Mountains, Colorado. National Technical Wildlife Diseases 29(1):118-122. Information Service, U.S. Department of Commerce, Wood, P. 1984. The elk hunt goes on at Grand Teton. Washington, D.C. National Parks 58(9-10):29-31. MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 55

Appendix A.

Duration of monitoring of 97 radio-collared elk (Cervus elaphus) of the Jackson elk herd, Wyoming, 1978-1984. Duration of monitoring (months) <12 >24 Monitoring interval (days) - Year N Range X SE % % Year X SE 1978 24 0-32 13.3 1.9 41.7 8.3 1978 5.6 0.4 1979 17 1-14 5.3 1.1 94.1 0 1979 5.6 0.4 1980 29 0-60 32.4 3.5 17.2 62.1 1980 6.2 0.8 1981 21 6-48 22.0 3.2 19.0 33.3 1981 7.8 0.7 1982 6 2-36 15.2 6.3 66.7 33.3 1982 5.9 0.7 1978-79 41 0-32 9.4 1.3 64.4 4.9 1983 4.9 0.4 1980-82 56 0-60 27.3 2.4 21.3 48.2 1984 10.0 3.0 1978-82 97 0-60 19.2 1.6 40.2 29.9 1978-84 6.5 0.6 56 RESOURCE PUBLICATION 199

Appendix B.

Dosages of powdered succinyl-choline chloride used to immobilize elk (Cervus elaphus) at the National Elk Refuge, Wyoming, 1978-1983, and the duration of immobilization and recovery times of the elk. Dosage Immobilization time Recovery time (mg) (min) {min2 xSD xSD xSD Sex N (Range) (Range) (Range) Male 19 22.9 ± 0.78 6.8 ± 2.85 38.4 ± 10.15 (22-24) (1-12) (22-60) Female 40 21.1 ± 0.93 9.4 ± 5.42 37.3 ± 11.49 (20-23) (4-25) (16-54) Total 59 21.7 ± 1.23 8.6 ± 4.84 37.7 ± 11.02 (20-24) (1-25) (16-60) Appendix C.

Hunting regulations for the Jackson elk (Cervus elaphus) herd by hunting area, 1978-1986.a Area 1986 1985 1984 Area 70 Buffalo Fork Seasons 10 Sep-31 Oct: Mature males only 10 Sep-31 Oct: Mature males only 10 Sep-2 Nov: Antlered and 50 any-elk permits, spikes excluded 50 any-elk permits Area 71 Pacific Creek Seasons 10 Sep-31 Oct: Mature males only 10 Sep-31 Oct: Mature males only 10 Sep-11 Nov: Antlered and 15 Oct-14 Nov: 100 any, spikes 50 any permits, spikes excluded 50 any permits excluded 27 Oct-11 Nov: 50 any permits Area 72 Berry Creek Closed Closed Closed Area 74 Ditch Creek Seasons Combined with Area 81 Combined with Area 81 Combined with Area 81 ~ Area 75 Riverbottom ~ Seasons 25 Oct-14 Nov: Any (permit only) 26 Oct-15 Nov: Any (permit only) 27 Oct-11 Nov: Any (permit only) Permits 90 each week 90 each week 90 each week U)~ Total Permits 270 270 270 Area 76 Blacktail/hayfields ~ Seasons 25 Oct-14 Nov: Any (permit only) 26 Oct-15 Nov: Any (permit only) 27 Oct-11 Nov: Any by permit Permits 160 a week of which 90 for 160 a week of which 90 for 160 a week of which 90 for only Blacktail Butte only Blacktail Butte only Blacktail Butte Total Permits 480 480 480 I Area 77 National Refuge Seasons ; 25 Oct-14 Nov: any (permit only) 26 Oct-15 Nov: Any (permit only) 27 Oct-16 Nov: Any by permit Permits 120 week, maximum of 40 120 each week, maximum of 40 120 each week, maximum of 40 ~ hunters at a time hunters at a time hunters at a time Total Permits 360 any 360 any 360 any i Area 78 Wilson Seasons Closed Closed 15 N ov-15 Dec: Archery only Area 79 Teton Park Seasons 25 Oct-14 Nov: Any (permit only) 26 Oct-15 Nov: Any (permit only) 27 Oct-11 Nov: Any (permit only) It_:i:j Permit Type 1 1,500 any 1,000 any 1,000 any Permit Type 2 None None 10 Nov-11 Nov: 800 any ~ Area 80 Sheep Creek E!1 Seasons 10 Sep-31 Oct: Mature males only 10 Sep-31 Oct: Mature males only 10 Sep-2 Nov: Antlered § Area 81 Spread Creek 01 -:t en Appendix C. Continued. 00 Area 1986 1985 1984 ~ Ul Seasons 10 Sep-31 Oct: Mature males only 10 Sep-31 Oct: Mature males only 10 Sep-2 Nov: Antlered & 100 antlerless permits & 100 any permits, spikes & 200 any permits 0~ excluded t:r:l Area 82 Crystal Creek Seasons 10 Sep-31 Oct: Mature males only 10 Sep-31 Oct: Mature males only 10 Sep-2 Nov: Antlered g ~ & 100 antlerless permits & 100 any permits, spikes & 200 any permits 0 excluded ~ Area 83 Fish Creek Seasons 1 Oct-31 Oct: Mature males only 1 Oct-31 Oct: Mature males only 1 Oct-31 Oct: Antlered .....~ & 100 antlerless permits & 100 any permits, spikes & 200 any permits "' excluded "' Area 1983 1982 1981

Area 70 Buffalo Fork Seasons 10 Sep-15 Nov: Antlered 10 Sep-29 Oct: Antlered 10 Sep-30 Oct: Antlered 29 Oct-4 Nov: Any 30 Oct-15 Nov: Any 31 Oct-15 Nov: Any 16 N ov-06 Dec: Any in part of area Area 71 Pacific Creek Seasons 10 Sep-28 Oct: Antlered 10 Sep-29 Oct: Antlered 10 Sep-30 Oct: Antlered 29 Oct-30 Nov: Any 30 Oct-31 Dec: Any 31 Oct-31 Dec: Any Area 72 Berry Creek Closed Closed Closed Area 7 4 Ditch Creek Seasons 10 Sep-15 Nov: Antlered 10 Sep-24 Oct: antlered 10 Sep-20 Oct: Antlered 24 Oct-04 Nov: Any 25 Oct-05 Dec: Antlerless 21 Oct-06 Dec: Antlerless Area 75 Riverbottom Seasons 29 Oct-30 Nov: 33 antlerless 30 Oct-05 Dec: 54 antlerless 31 Oct-06 Dec: 30 antlerless & 12 any permits at a time & 6 any permits at a time & 6 any permits at a time Permits 45 permits per time period 60 permits per time period 36 permits per time period 170 any; 467 anterless 95 any; 539 antlerless 95 any; 476 antlerless Area 76 Blacktail/hayfields Seasons 29 Oct-30 Nov: Antlerless by 30 Oct-05 Dec: Antlerless by 31 Oct-06 Dec: Antlerless by permit only permit only permit only Permits 135 a period (18 any) of 150 a period (18 any) of 72 a period (6 any) of which 60 are Blacktail Butte which 60 are Blacktail Butte which 36 are Blacktail Butte Total Permits 255 any; 1,655 antlerless 95 any; 2,283 antlerless 95 any; 1,047 antlerless Area 77 National Elk Refuge Seasons 29 Oct-30 Nov: 10 any & 30 30 Oct-05 Dec: Any until 31 Oct-06 Dec: Any until Appendix C. Continued. -- Area 1983 1982 1981 antlerless by permit only migration, then antlerless only migration, then antlerless only Permits 120 a week, maximum 40 a day 150 a week, maximum 50 a day 120 a week, maximum 40 a day Total Permits 320 any; 44 7 antlerless 385 any; 646 antlerless 596 any; 530 antlerless Area 78 Wilson Seasons 15 Nov-15 Dec: Archery only 15 Nov-05 Dec: Archery only 15 N ov-15 Dec: Antlerless Area 79 Teton Park Seasons 29 Oct-30 Nov: Any (permit only) 30 Oct-05 Dec: Any (permit only) 31 Oct-06 Dec: Any (permit only) Permit Type 1 1,500 any 1,500 any 1,500 any Permit Type 2 12 Nov-30 Nov: 1,000 any 13 Nov-05 Dec: 1,500 any 14 Nov-06 Dec: 1,000 any Area 80 Sheep Creek Seasons 10 Sep-04 Nov: Antlered 10 Sep-24 Oct: Antlered 10 Sep-20 Oct: Antlered 25 Oct-10 Nov: Any 21 Oct-06 Nov: Any 11 Nov-05 Dec: Antlerless, 07 Nov-06 Dec: Antlerless, noon opening noon opening Area 81 Spread Creek ~ Seasons 10 Sep-31 Oct: Antlered 10 Sep-31 Oct: Antlered 10 Sep-20 Oct: Antlered 0 24 Oct-04 Nov: Any 25 Oct-15 Nov: Any 21 Oct-15 Nov: Any - 16 Nov-05 Dec: Antlerless 16 Nov-06 Dec: Antlerless ~ Area 82 Crystal Creek 0-z Seasons 10 Sep-31 Oct: Antlered 10 Sep-31 Oct: Antlered 10 Sep-31 Oct: Antlered en 10 Sep-15 Nov: 250 any permits 10 Sep-15 Nov: 300 any permits 01 Oct-15 Nov: 300 any permits t:,~ Area 83 Fish Creek Seasons 01 Oct-31 Oct: Antlered and 01 Oct-31 Oct: Antlered and 01 Oct-31 Oct: Antlered and 300 any permits 350 any permits 300 any permits I0 Area 1980 1979 1978 t_,:j ~ ~ Area 70 Buffalo Fork 0 Seasons 20 Sep-17 Oct: Antlered 10 Sep-19 Oct: Antlered 10 Sep-20 Oct: Antlered >'%j >-,3 18 Oct-15 Nov: Any 20 Oct-15 Nov: Any 21 Oct-15 Nov: Any ::c: 16 Nov-07 Dec: Any inl-9 Dec: Anyl-15 Dec: Any t_,:j

part of area (j~ Area 71 Pacific Creek @ Seasons 10 Sep-17 Oct: Antlered 10 Sep-19 Oct: Antlered 10 Sep-20 Oct: Antlered z0 18 Oct-31 Dec: Any 20 Oct-31 Dec: Any 21 Oct-31 Dec: Any t,:j Area 72 Berry Creek ~ Closed Closed Closed ::c: Area 74 Ditch Creek t_,:j ~ Seasons 10 Sep-17 Oct: Antlered 10 Sep-19 Oct: Antlered 10 Sep-20 Oct: Antlered t:,

18 Oct-17 Dec: Antlerless 20 Oct-09 Dec: Antlerless 21 Oct-03 Dec: Antlerless 01 ~ Appendix C. Continued. O') 0 Area 1980 1979 1978 Area 75 Riverbotton ~rn Seasons 25 Oct-07 Dec: 25 antlerless 27 Oct-09 Dec: Antlerless 28 Oct-03 Dec: Antlerless & 5 any permits at a time by permit only Permits 30 permits per time period 20 permits per week 20 permits per week i Total Permits 95 any; 4 71 antlerless 251 106 Area 76 Blacktail/hayfields Seasons 25 Oct-07 Dec: Antlerless 28 Oct-03 Dec: Antlerless 27 Oct-09 Dec: Antlerless i0 by permit only by permit only by permit only ~ Permits 70 a period (10 any) of 60 a week; 40 for hayfields; 60 a week; 40 for hayfields; Total Permits ~ which 25 are Blacktail Butte 20 for Blacktail Butte 20 for Blacktail Butte ...... 190 any; 1,131 antlerless 503 317 ~ Area 77 National Refuge ~ Seasons 25 Oct-07 Dec: any until 20 Oct-02 Nov: Any 28 Oct-08 Dec: Antlerless by migration; then only 03 Nov-09 Dec: Antlerless permit only antlerless by permit only Permits 120 a week; maximum of 40 a day 120 a week; maximum of 40 a day 120 a week; maximum of 40 a day Total Permits 1,269 any; 280 Antlerless 246 any; 778 Antlerless 639 Antlerless Area 78 Wilson Seasons 15 Nov-15 Dec: Antlerless 15 N ov-15 Dec: Antlerless 15 Nov-15 Dec: Antlerless Area 79 Teton Park Seasons 25 Oct-07 Dec: Any by permit 27 Oct-09 Dec: Any by permit 28 Oct-03 Dec: Any by permit Permit Type 1 l,500Any l,500Any l,500Any Permit Type 2 08 Nov-07 Dec: 1,000 Any 10 Nov-09 Dec: 1,000 Any 11 Nov-03 Dec: 1,000 Any Area 80 Sheep Creek Seasons 10 Sep-17 Oct: Antlered 10 Sep-19 Oct: Antlered 10 Sep-27 Oct: Antlered 18 Oct-31 Oct: Any 20 Oct-02 Nov: Any 28 Oct-08 Dec: Antlerless, 01 Nov-07 Dec: Antlerless, 03 Nov-09 Dec: Antlerless, noon opening noon opening noon opening Area 81 Spread Creek Seasons 10 Sep-17 Oct: Antlered 10 Sep-19 Oct: Antlered 10 Sep-20 Oct: Antlered 18 Oct-15 Nov: Any 20 Oct-15 Nov: Any 21 Oct-15 Nov: Any 16 Nov-07 Dec: Antlerless 16 N ov-09 Dec: Antlerless 16 N ov-03 Dec: Antlerless in part of area Area 82 Crystal Creek Seasons 10 Sep-15 Nov: Antlered 10 Sep-19 Oct: Antlered 10 Sep-20 Oct: Antlered 18 Oct-15 Nov: 300 Any permits 20 Oct-15 Nov: 300 Any permits 21 Oct-15 Nov: 300 Any permits Area 83 Fish Creek Seasons 01 Oct-31 Oct: Antlered 01 Oct-14 Oct: Antlered 01 Oct-14 Oct: Antlered 18 Oct-15 Nov: 300 Any permits 15 Oct-31 Oct: 250 Any permits 15 Oct-31 Oct: 250 Any permits a see Fig. 3. MIGRATIONS AND MANAGEMENT OF THE JACKSON ELK HERD 61

Appendix D.

Cattle allotments in the Bridger-Teton National Forest and Grand Teton National Park. Allotmenta Number of cattle Date inc Date ouf Bridger-Teton National Forest A-Pacific Creek 249 6/1 8/22 B-Lava Creek East Unit 271 6/15 10/15 C-Lava Creek West Unitb D-Burro Hill 50 6/15 10/15 E-Fish Creek 1,959 6/11 10/15 F-Upper Gros Ventre 750 6/18 10/8 G-Redmond Creek 30 6/15 9/26 H-Taylor cattle and horse 15 5/15 11/15 I-Kinky Creek 174 6/11 10/15 J-Blackrock-Spread Creek 1,456 6/15 10/15 K-Ditch Creek 350 7/1 10/31 L-Miner's Creek 92 6/21 10/15 M-Bacon Creek 1,677 6/11 10/15 N-Big Cow Creek-Robinson 32 5/16 11/30 0-Bierer Creek 30 6/15 9/26 P-Breakneck 750 6/18 10/8 Grand Teton National Park Q-Gros Ventre pasture 1,200 5/15 6/15 R-Cunningham pasture 975 6/15 7/10 975 10/15 11/15 S-Elk Ranch West 975 6/15 7/10 975 10/15 11/15 T-Elk Ranch East 975 7/10 10/14 U-Sread Creek-Uhl 299 5/29 6/15 V-Pacific Creek 150 6/10 9/15 a Figure 11 shows the location of the lettered allotments in the Jackson elk herd unit. bused in alternate years with Lava Creek East and Burro Hill. c Grazing dates. A list of current Resource Publications follows.

177. Field Manual for the Investigation of Fish Kills, by Fred P. Meyer and Lee A. Barclay, editors. 1990. 120 pp. 178. Section 404 and Alterations in the Platte River Basin of Colorado, by Douglas N. Gladwin, Mary E. Jennings, James E. Roelle, and Duane A. Asherin. 1992. 19 pp. 179. Hydrolgy of the Middle Rio Grande From Velarde to Elephant Butte Reservoir, New Mexico, by Thomas F. Bullard and Stephen G. Wells. 1992. 51 pp. 180. Waterfowl Production on the Woodworth Station in South-central North Dakota, 1965-1981, by Kenneth F. Higgins, Leo M. Kirsch, Albert T. Klett, and Harvey W. Miller. 1992. 79 pp. 181. Trends and Management of Wolf-Livestock Conflicts in Minnesota, by Steven H. Fritts, William J. Paul, L. David Mexch, and David P. Scott. 1992. 27 pp. 182. Selection of Prey by Walleyes in the Ohio Waters of the Central Basin of Lake Erie, 1985-1987, by David R. Wolfert and Michael T. Burr. 1992. 14 pp. 183. Effects of the Lampricide 3-Trifluoromethyl-4-Nitrophenol on the Pink Heelsplitter, by Terry D. Bills, Jeffrey J. Rach, LeifL. Marking, and George E. Howe. 1992. 7 pp. 184. Methods for Detoxifying the Lampricide 3-Trifluoromethyl-4-Nitrophenol in a Stream, by Philip A. Gilderhus, Terry D. Bills, and David A. Johnson. 1992. 5 pp. 185. Group Decision-making Techniques for Natural Resource Management Applications, by Beth A. Coughlan and Carl L. Armour. 1992. 55 pp. 186. DUCKDATA: A Bibliographic Data Base for North American Waterfowl (Anatidae) and Their Wetland Habitats, by Kenneth J. Reinecke and Don Delnicki. 1992. 7 pp. 187. Dusky Canada Goose: An Annotated Bibliography, by Bruce H. Campbell and John E. Cornely. 1992. 30 pp. 188. Human Disturbances of Waterfowl: An Annotated Bibliography, by Robert B. Dahlgren and Carl E. Korshgen. 1992. 62 pp. 189. Opportunities to Protect lnstream Flows and Wetland Uses of Water in Nevada, by James L. Bingham and George A. Gould. 1992. 33 pp. 190. Assessment of Habitat of Wildlife Communities on the Snake River, Jackson, Wyoming, by Richard L. Schroeder and Arthur W. Allen. 1992. 21 pp. 191. Evaluating Temperature Regimes for Protection of Smallmouth Bass, by Carl L. Armour. 1993. 26 pp. 192. Sensitivity of Juvenile Striped Bass to Chemicals Used in Aquaculture, by Terry D. Bills, Leif L. Marking, and George E. Howe. 1993. 11 pp. 193. Introduction of Foxes to Alaskan Islands-History, Effects on Avifauna, and Eradication, by Edgar P. Bailey. 1993. 53 pp. 194. Distribution and Abundance of Predators that Affect Duck Production-Prairie Pothole Region, by Alan B. Sargeant, Raymond J. Greenwood, Marsha A. Sovada, and Terry L. Shaffer. 1993. 96 pp. 195. Evaluating Temperature Regimes for Protection of Walleye, by Carl L. Armour. 1993. 22pp. 196. Evaluation of Five Anesthetics on Striped Bass, by Carol A. Lemm. 1993. 10 pp. 197. Standardization of Roadside Counts of Columbids in Puerto Rico and on Vieques Island, by Frank F. Rivera-Milan. 1993. 26 pp. 198. Herpetofaunal Diversity of the Four Holes Swamp, South Carolina, by Russell J. Hall. 1994. 43 pp. U.S. DEPARTMENT OF THE INTERIOR NATIONAL BIOLOGICAL SURVEY

As the Nation's principal conservation agency, the Department of the Interior has responsibility for most of our nationally-owned public lands and natural resources. This includes fostering the sound use of our lands and water resources; protecting our fish, wildlife, and biological diversity; preserving the environmental and cultural values of our national parks and historical places; and providing for the enjoyment of life through outdoor recreation. The Department assesses our energy and mineral resources and works to ensure that their development is in the best interests of all our people by encouraging stewardship and citizen participation in their care. The Department also has a major responsibility for American Indian reservation communities and for people who live in island territories under U.S. administration.